昆虫的夏滞育

夏滞育作为昆虫越夏的生态策略 ,已广泛引起昆虫学家的重视。本文就几十年来夏滞育研究的进展进行综述。着重讨论诱导、维持、终止夏滞育的主要环境因子 ,夏滞育的生态特征和它的生态适应意义。     

昆虫夏滞育的调控及其遗传基础

几十年来对昆虫夏滞育的研究不断深入 ,阐明了不少昆虫夏滞育的生态机理。目前 ,探讨其内在的机制已成为研究的方向。该文着重论述夏滞育的生理生化特征、夏滞育的调控及其遗传基础     

Monitoring diapause development in the Colorado potato beetle, Leptinotarsa decemlineata, under field conditions using molecular biomarkers

A multiplex PCR protocol was developed using five diapause-regulated genes to monitor diapause development of the Colorado potato beetle under field conditions. A total of 870 beetles from the Red River valley of North Dakota and Minnesota, USA, were screened for three consecutive years. Out of the 32 possible expression profiles, eight could be arranged in chronological order of diapause development. These eight profiles account for over 92% of the beetles surveyed. Intra-population variation in diapause phenotypes was observed in the field. Some beetles were already in the diapause initiation phase in June when the day length was greater than 17 h. Inter-seasonal variation in the timing of diapause development was also noted. The greatest differences were before the day length decreased to less than 15 h. Anomalies in the results, e.g., the presence of the diapause maintenance phase profiles in beetles collected on the potato plants, argue that laboratory results are not always equivalent with what is observed under field conditions.     

Photoperiodic control of adult diapause in the predacious mite, Amblyseius potentillae: repeated diapause induction and termination

ABSTRACT. Photoperiodic control of facultative reproductive diapause was examined in females of the phytoseiid mite, Amblyseius potentillae (Garman). Full diapause was induced by rearing the mites from egg to adult under short-day photoperiods. Egg-laying females, however, which had experienced a long-day photoperiodic regime during pre-imaginal development, could still be induced to enter diapause when they were transferred to a short-day regime. Diapause development proceeded slowly under a short-day photoperiod, but was accelerated under long days. During diapause development under short days the photoperiodic sensitivity of the females diminished gradually and ultimately disappeared completely. However, after completion of diapause, sensitivity to photo-period reappeared. A second diapause could be induced in post-diapause females under short-day photoperiods and completed again under long-day photoperiods. These results show that A. potentillae remains sensitive to diapause-inducing and diapause-averting daylengths during the adult stage and that a second diapause may be induced after completion of the first one.     

Effect of temperature and photoperiod on diapause development in a Douglas fir seed chalcid, Megastigmus spermotrophus

Douglas fir seeds infested by diapausing larvae of a chalcid, Megastigmus spermotrophus, were transferred under different temperature and photoperiod regimes at various dates during autumn and winter. Both winter diapause and prolonged diapause were compared to the diapause patterns observed in control lots kept under natural conditions. The results showed that exposure to low temperatures in autumn and winter is required for the completion of diapause development. Median adult emergence dates and the date of transfer of larvae to 20°C were negatively correlated. The frequency of prolonged diapause of larvae transferred to 20°C decreased significantly after November. When larvae were submitted to different constant temperatures, the longer the daylength, the higher and faster the reactivation. These results suggest that chalcid larvae can perceive daylength through the seed coat. This phenomenon is probably related to the fact that most of the infested seeds falling in autumn are usually covered by a layer of leaf litter during the winter. Received: 14 June 1996 / Accepted: 24 February 1997     

The roles of dispersal limitation and environmental conditions in controlling caddisfly (Trichoptera) assemblages

1. Many recent studies have quantified the relative importance of environmental variables and dispersal limitations in shaping the structure of stream communities. However, the relative importance of these factors at different spatial extents has been seldom evaluated. 2. We analysed the distribution of caddisfly species in 89 Amazonian streams in relation to stream characteristics and spatial variables representing overland dispersal routes. The streams occur in three regions that differ in spatial extent and environmental characteristics. We analysed the data using partial redundancy analysis with two predictor data sets, one environmental and one spatial, to evaluate the variation in assemblage composition. We also separated caddisflies into ‘good’ and ‘poor’ dispersers to evaluate possible differences in the responses of these two groups. 3. The environmental component explained a higher proportion of variance in assemblage composition than did the spatial component. Spatial effects were evident only when data from all three regions were analysed together, although the exclusive spatial fraction was quite low. Good dispersers responded similarly to the community as a whole, while poor dispersers were related to environmental variables only in one region and also were not related to spatial variables. 4. Caddisflies were most affected by environmental factors. The large environmental effect and small spatial effect are in accord with the use of these stream insects as good indicators of site properties and disturbances in monitoring programmes.     

松毛虫赤眼蜂滞育诱导及解除条件研究

【目的】以柞蚕Antheraea pernyi卵为繁殖寄主,对松毛虫赤眼蜂Trichogramma dendrolim滞育诱导及解除条件进行研究,以解决赤眼蜂工厂化生产和大面积应用中面临的的中、长期储存问题。【方法】通过观测不同发育阶段(寄生柞蚕卵在26℃培养40、96和144 h)、滞育诱导温度(10、13和16℃)和诱导时间对松毛虫赤眼蜂滞育的影响,确定松毛虫赤眼蜂滞育诱导条件;通过观测滞育诱导温度和滞育后的贮藏温度对滞育解除的影响,确定松毛虫赤眼蜂滞育解除条件。【结果】在松毛虫赤眼蜂的不同发育阶段对其进行持续的低温刺激均能使其导入滞育,但以小幼阶段(26℃培养40 h)开始效果最佳,寄生卵在26℃培养40 h后,转入10℃和13℃下连续诱导31 d,滞育率可达100%和99.12%。滞育诱导温度和滞育后的贮藏温度对松毛虫赤眼蜂解除滞育所需时间和解除滞育后的羽化出蜂率有较大影响,10℃诱导滞育后置于1℃冷藏的赤眼蜂解除滞育所需时间最短,解除滞育后的羽化出蜂率和单卵出蜂数更高,更耐储存。此条件下冷藏约30 d开始打破滞育,在正常发育下温度下羽化出蜂,60 d羽化出蜂率达到95.24%,冷藏4个月后羽化出蜂率仍在60%以上,单卵出蜂数高于50头。【结论】松毛虫赤眼蜂最佳滞育诱导条件为26℃培养40 h后,转入10℃连续低温诱导31 d;最佳滞育解除条件为1℃低温储存,但储存期不能超过4个月。     

Negative genetic correlation between diapause duration and fecundity after diapause in a spider mite

Abstract.  1. Recently, the costs of diapause, i.e. the reduction of fecundity after diapause, have been examined from an evolutionary perspective.
2. The evolution of this trade-off should be clarified by quantitative genetic approaches, as theoretical studies address the evolution of multiple traits. Nevertheless, previous studies on the costs of diapause have been based on phenotypic correlations or experimental manipulations, whereas the genetic background underlying this trade-off remains unclear.
3. In the present study, a half-sib breeding design was used to examine the quantitative genetic relationships between diapause duration and post-diapause fecundity in the Kanzawa spider mite Tetranychus kanzawai Kishida (Acari: Tetranychidae).
4. The heritability of diapause duration, post-diapause total fecundity, and post-diapause early fecundity were 0.37, 0.14, and 0.11 respectively. Genetic correlations between diapause duration and post-diapause total fecundity, and between diapause duration and early fecundity were both significantly negative (–0.70 and –0.90 respectively). These results suggest that the cost of prolonging diapause duration is genetically based, and that these life-history traits respond to natural selection acting on them simultaneously.     

Geographic variation in photoperiodic diapause induction and diapause intensity in Sericinus montelus (Lepidoptera: Papilionidae)

Abstract Due to the risk of extinction and ornamental value of the swallowtail butterfly, Sericinus montelus Gray (Lepidoptera: Papilionidae) in China, knowledge about local adaptations is important for the conservation and economical utilization of the species. In the present study, photoperiodic diapause induction and diapause intensity of S. montelus populations from Jiamusi (46°37′N), Beijing (40°15′N), Zibo (36°48′N), Fangxian (32°36′N), Wuhan (30°33′N) and Huaihua (27°33′N) were characterized at 25°C. Logistic regression analysis revealed a significant population × hours of light interaction, confirming that photoperiodic responses varied among populations. The critical photoperiod was positively correlated with latitude and increased toward the north at a rate of about 1 h for each 6.67 degrees of latitude. Survival analyses indicated that survival time of diapausing pupae before adult eclosion differed significantly among populations at 25°C and 16 : 8 L : D h. The mean duration of pupal diapause was also positively correlated with latitude. Our study reveals geographic variation in the critical photoperiod for diapause induction and in diapause intensity of S. montelus. These results provide useful information for our general understanding about seasonal adaptation in insects and may also be used to predict how geographic populations respond to climate warming.     

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.     

Glycerol and glucose accumulation during diapause in a tropical beetle

Abstract. Carbohydrate changes were measured during the 10-month adult diapause in the neotropical beetle, Stenotarsus rotundus Arrow (Endomychidae, Coleoptera) to test the hypothesis that metabolic suppression causes the accumulation of polyols in the absence of the need for cold tolerance. Glycerol and glucose both accumulate during the first 3 months from June to September, but decline in October, accumulate again in December-January and finally decline after January and stay low until the adults leave their diapause site in April. Any adaptive significance for this pattern is unknown but its coincidence with previously measured metabolic suppression suggests that glycerol and glucose accumulate as a result of metabolic adjustments during diapause. The relevance of these results to the evolution of polyol accumulation for cryoprotection in temperate insects in discussed.     

Thermal response and reversibility of diapause in the eggs of Locusta migratoria

Abstract. Eggs of a local population of Locusta migratoria L. (Orthoptera: Acrididae) collected near Hirosaki (40.5^oN) entered diapause when incubated at temperatures between 20 and 35^oC. For diapause development the optimum temperature was 10^oC. The lower thermal threshold for post-diapause development was 14.7^oC. After chilling at 10^oC for 20 days, the rate of hatching varied with incubation temperature, being 0, 61% and 81% at 20, 25 and 30^oC, respectively. After chilling for 40 days or more, however, almost all eggs hatched at 20–30^oC. Diapause with a reduced intensity seemed to be eliminated easily by a high temperature of 25 or 30^oC.
When eggs chilled at 10^oC for 20 days were kept at 20^oC for 7 days or more before incubation at 25^oC, almost all eggs maintained diapause. Most eggs chilled at 10^oC for three 10 day periods separated by 3 days of warming at 25^oC failed to terminate diapause. Daily alternations of 10^oC (18 h) and 25^oC (6h) decreased the diapause-terminating effect of chilling. These facts suggest that diapause intensity can be restored if eggs chilled insufficiently are exposed to a moderately high temperature. This reversible change in diapause intensity would play an important role in maintaining diapause before winter.     

Effect of daylength and temperature on the larval diapause of the sunflower moth, Homoeosoma electellum

The effect of daylength and temperature on the regulation of the larval diapause of a central Missouri population of the sunflower moth, Homoeosoma electellum, was examined. Fully grown fourth-instar larvae exhibit a facultative diapause. Measurements of the effect of photoperiod on diapause induction revealed critical photoperiods of about 13 h 30 min light/day at 20°C, and between 11 h 45 min and 12 h light/day at 23°C. Third and fourth-instar larvae were shown to be the main sensitive stages for diapause determination. Daylength was also shown to be an important regulator of the rate of diapause development. A short day of LD 10:14 h permitted only a low rate of diapause development, whereas long days of LD 14:10 h and LD 16:8 h accelerated diapause development at 25 and 30°C. When long days were alternated with short days at 30°C the accelerating effect of long days on diapause development was not found. Systematic transfers of chilled diapausing larvae revealed an accelerated diapause development in groups transferred from 10 to 30°C LD 10:14 h, but diapause development was not accelerated in groups transferred from 10 to 30°C LD 16:8 h.     

Geographic variation in diapause induction and mode of diapause inheritance in Tetranychus pueraricola

Abstract:  Diapause induction and photoperiodic response curves were determined for 33 strains of Tetranychus pueraricola derived from kudzu vine at three constant temperatures (15, 18 and 20°C) under a short-day condition (10 : 14 h; light : dark). Females of all but one of the strains entered diapause at all three temperatures with little variation in diapause percentages among the strains. The exception was the southernmost strain, which was found to be a non-diapause (ND) strain. The critical photoperiod gradually decreased towards the south at a rate of about 1 h for each 5 degrees of latitude. The diapause strains (D1 and D2) exhibited 100% diapause, whereas the ND strain exhibited 0% diapause. By crossing these strains, we determined that 'non-diapause' was a dominant character over 'diapause' and the character was controlled by simple Mendelian inheritance. To clarify why the female progeny from the crosses between the D1 and ND strains did not segregate into the diapause and non-diapause phenotypes in a 1:1 ratio in the B₁ generation, round-robin crosses were carried out among the three strains. The results showed that the F₁ generation was reproductively compatible and showed high egg hatchability with a female-biased sex ratio. In the B₁ generation, the crosses between the D1 and ND strains and between the D1 and D2 strains exhibited extremely low egg hatchability and produced mostly female progeny, whereas offspring from the crosses between the D2 and ND strains showed more than 50% hatchability for B₁ eggs and a female-biased sex ratio. Thus, the absence of segregation observed in the crosses between the D1 and ND strains appears to be due to the severe hybrid breakdown that occurred in the B₁ generation.     

Significance of water contact as a factor terminating larval diapause in a stem borer, Busseola fusca

Previous studies revealed that diapause larvae of Busseola fusca (Lepidoptera: Noctuidae) began to respond to water and pupated in artificial wet conditions only in late diapause. Experiments are designed to clarify how water is involved in the termination of larval diapause. Neither feeding the larvae on fresh sorghum stems (6 weeks old) nor allowing them to drink water stimulated a break in the larval diapause. Diapause larvae transferred to artificial wet conditions pupated without increase in fresh weight. It is thus concluded that water contact is more significant as a factor terminating diapause in B. fusca than water uptake.

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Résumé Des études antérieures avaient montré que la diapause des chenilles de B. fusca (Lep., Noctuidae) commence par une réaction à l'eau et qu'elles ne se nymphosent dans des conditions artificiellement humides qu'en fin de diapause. Des expériences ont été conçues pour préciser l'intervention de l'eau dans la terminaison de la diapause. Ni l'alimentation des chenilles sur des tiges fraiches de jeune sorgho (de 6 semaines), ni la possibilité de boire de l'eau n'ont stimulé une rupture de la diapause larvaire. Des chenilles en diapause transférées dans des conditions artificiellement humides se sont nymphosées sans augmentation du poids frais. On en a conclu que le contact avec de l'eau était plus important que l'absorption d'eau comme facteur de fin de diapause chez B. fusca.

     

Photoperiodic induction of larval diapause and temperature-dependent termination in a wild multivoltine bruchid,Kytorhinus sharpianus

A wild bean weevil,Kytorhinus sharpianus Bridwell (Coleoptera: Bruchidae), has a multivoltine life cycle and enters a hibernal larval diapause at the fourth instar under a short daylength (Shimada & Ishihara, 1991). Here, we investigated their diapause incidence under different photoperiods at 24°C and 27°C. The critical photoperiods for diapause induction were 14.5 h at 24°C and 14 h at 27°C. The stages susceptible to diapause-inducing stimuli were estimated by transferring larvae of various instars from long days to short days and vice versa. Then we investigated the incidence of larval diapause. The sensitive stage was estimated to be from the third to early fourth instar. Though larval diapause, which was induced under a short daylength, was terminated only by increasing the daylength, the termination was more synchronized by an exposure to a low temperature followed by increasing temperature, irrespective of photoperiod.     

大猿叶虫滞育进程中成虫内生殖系统的形态特征

大猿叶虫Colaphellus bowringi Baly是十字花科蔬菜的一种重要害虫,以成虫在土壤中滞育越冬和越夏。本研究通过解剖大猿叶虫非滞育成虫,观察描述了雌雄成虫内生殖系统的结构特点,并绘制了雌雄成虫内生殖系统模式图。对滞育初期、滞育期间、滞育解除后未取食和滞育解除后取食成虫解剖显示,滞育初期,雌雄成虫内生殖系统几乎与羽化初期成虫一致。滞育期间雄成虫的附腺和射精管亦不发达,滞育解除后未取食雄成虫的附腺膨大且粗于输精管,射精管略膨大,取食雄成虫的射精管呈不透明淡桔黄色。滞育期间雌成虫的卵巢小,略大于滞育初期卵巢,少数卵巢小管的基部可见具卵黄原沉淀的卵粒。滞育解除后未供食雌成虫卵巢明显膨大可见大量成形卵粒,部分卵粒可见卵黄原沉淀;取食雌成虫的卵巢膨大,可见大量成熟卵粒,侧输卵管和总输卵管中可见待产的卵。作者认为,大猿叶虫成虫在滞育期间能够缓慢发育,部分滞育前积累的代谢物质被用于滞育后发育,但只有经过取食,成虫才能正常交配和产卵。     

Repeated larval diapause and diapause-free development in geographic strains of the burnet moth Zygaena trifolii Esp. (Insecta,Lepidoptera)

Summary Zygaena trifolii is a long-day insect with temperature-dependent photoperiodic responses. All larval instars are sensitive to photoperiod; however, diapause may occur at the third larval stage or any subsequent larval instars. There were quantitative differences within populations in the threshold photoperiod for diapause induction. The diapause response was polymorphic, so that larvae might enter diapause at different instars under the same culture conditions. Furthermore, decreasing photoperiods below a critical daylength shifted the diapausing instar towards earlier stages. Geographic strains of Z. trifolii showed discontinuous clinal variation. Near the northern edge of the distribution [Cologne (Köln), FRG], there is first an obligatory diapause, mainly during early instars, and additional facultative (repeat) diapauses during later larval instars in subsequent years. In the southern part of its distribution, this burnet moth is partially bivoltine in the field with a facultative first developmental arrest and a decreased capacity for repeated diapause (Valencia, Spain; Marseille, France). Further experiments indicated that the photoperiodically controlled diapause reaction is also influenced by the number of photoperiodic cycles experienced during the period spent in each larval instar, which depends on temperature. The adaptive significance of obligatory and facultative repeated diapause, varying even among the offspring of a single female, may be to buffer the populations against the more extreme and, from year to year, unpredictable fluctuations in climatic conditions at the northern edge of the distribution.Abbreviations L₃ feeding 3rd larval instar - L₄D diapausing 4th larval instar - L₅D2 repeat-diapausing larval instar with second diapause at the 5th larval stage - LD light-dark cycle - KT shortday conditions (e.g. LD 8:16) - LT long-day conditions (e.g. LD 16:8)     

Initial diapause and embryonic development in the speckled bush-cricket, Leptophyes punctatissima

ABSTRACT. The initiation and pattern of embryonic development in Leptophyes punctatissima (Bosc) (Tettigoniidae) are shown to be variously dependent on temperature. Immediate high temperature (30°C) facilitated rapid and direct embryogenesis to the stage of diapause in the fully formed embryo. Slightly lower temperature (25°C) resulted in a delay before embryogenesis started, and this delay was greatly extended as an initial diapause if incubation temperatures were decreased (20-16°C). Still lower initial temperatures (8–12°C) facilitated subsequent development at 20–30°C. These responses all increased as a function of exposure time. Once initiated, the rate of development was temperature dependent, but competence to tolerate high temperature, and ability to continue development at low temperatures, changed with age. In general, the developmental temperature range appears to be lowered with age. None of these different treatments had any effect on the late embryonic diapause. According to the temperatures prevailing at oviposition, Leptophyes could be an annual or a biennial species; the biennial pattern is normal.