昆虫滞育研究进展

首先对昆虫滞育的若干名词概念作出解释,按照滞育过程的阶段特点划分为3个时期:(1)滞育前期(包括滞育诱导和滞育准备);(2)滞育期(包括滞育进入、滞育维持和滞育解除);(3)滞育后期。然后重点介绍这3个阶段近期分子和细胞水平上的重要工作,对一些科学问题进行讨论。     

Differences in the pre‐diapause and pre‐oviposition accumulation of critical nutrients in adult females of the beetle Colaphellus bowringi

In many insect species, the differentiation of development between diapause and reproduction first becomes obvious during the diapause preparation (pre‐diapause) and pre‐oviposition phases. However, the differentiation of nutrient accumulation between these two phases remains unclear. We compared the weights of pre‐diapause and reproductive adult female Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), and measured their triacylglycerol (TAG), protein, and carbohydrate content from emergence until they had fed for 4 days post‐eclosion. We also compared the ovarian development and accumulation of lipid droplets between pre‐diapause and reproductive adult females in order to determine whether we could visually detect differences in nutrient allocation. The weights of both pre‐diapause and reproductive females increased with duration of feeding. The fresh weight and water content of pre‐diapause females was significantly lower than that of reproductive females after feeding for 3 days post‐eclosion. Pre‐diapause females channeled their reserves into TAG in the fat body, whereas reproductive females converted nutrients into proteins and carbohydrates for egg development. These results quantify differences in nutrient accumulation between pre‐diapause and reproductive adult female C. bowringi, and provide clues for understanding the molecular mechanisms underlying differences in the allocation of nutrients between diapause and reproduction in insects.     

Effects of temperature and photoperiod on termination of pseudopupal diapause in the bean blister beetle, Epicauta gorhami

Larvae of the bean blister beetle, Epicauta gorhami (Coleoptera: Meloidae), feed on grasshopper eggs in soil and undergo hypermetamorphosis. This beetle undergoes larval diapause in the fifth instar as a pseudopupa, a form characteristic of hypermetamorphosis in meloid beetles. This pseudopupal diapause is maintained for a long period if the larvae are kept under continuous diapause-inducing short-day conditions. In the present study, the effects of temperature and photoperiod on termination of pseudopupal diapause in E. gorhami were examined using pseudopupae obtained under 12L-12D at 25°C. Diapause was terminated by exposure to low or high temperature, but not by transfer to long-day conditions. The pseudopupal stage comprises diapause and post-diapause phases despite its unchanging appearance. The optimum low temperature for diapause termination was 10-15°C, and a higher or lower temperature was less effective. There was an individual variation in the low temperature requirement for diapause termination and post-diapause development may not occur until this requirement is satisfied. Although under natural conditions pseudopupae may encounter low temperatures that are effective for diapause termination at different ages, our results show that pseudopupae at various ages respond similarly to low temperature exposure. This response may ensure that resumption of development synchronizes in a population.     

Termination of pupal diapause in the pine processionary moth Thaumetopoea pityocampa

Diapause development is a complex process involving several eco‐physiological phases. Understanding these phases, especially diapause termination, is vital for interpreting the life history of many insect species and for developing suitable predictive models of population dynamics. The pine processionary moth is a major defoliator of pine and a vertebrate health hazard in the Mediterranean region. This species can display either univoltine or semivoltine development, with a pupal diapause extending from a few months to several years, respectively. Although the ecological and applied importance of diapause is acknowledged, its physiological regulation in either case remains obscure. In the present study, we characterize pre‐termination, termination and post‐termination phases of pupae developing as univoltine or remaining in prolonged diapause. Changes in metabolic activity are monitored continuously using thermocouples, comprising a novel method based on direct calorimetry, and periodically by use of O₂ respirometry. The two methods clearly detect diapause termination in both types of pupae before any visible morphological or behavioural changes can be observed. Univoltine individuals are characterized by an increase in metabolic activity from pre‐termination through to termination and post‐termination, ultimately resulting in emergence. Remarkably, a synchronous termination is observed in individuals that enter prolonged diapause instead of emerging; however, in these pupae, the increased metabolic activity is only transient. The present study represents a starting point toward understanding the eco‐physiology of diapause development processes in the pupae of the pine processionary moth.     

Diapause development and acclimation regulating enzymes associated with glycerol synthesis in the Shonai ecotype of the rice stem borer larva,Chilo suppressalis walker

Overwintering larvae of the Shonai ecotype of the rice stem borer, Chilo suppressalis, enter diapause in early September and terminate diapause at the end of October. Cold acclimation at 0°C did not influence glycerol, trehalose or glycogen content in larvae collected on 22 September. Acclimation at 0°C increased the glycerol content and reduced the glycogen content significantly in larvae collected on 2 October and 22 November compared with acclimation at 15°C. These results indicate that overwintering larvae at different phases of diapause development respond differently to the low temperature stimulus for glycerol synthesis. Thus, we evaluated the metabolic rearrangements associated with glycerol synthesis during diapause development and after temperature acclimation. Larvae collected on 2 October were acclimated at 15°C for 15 and 60 days. Some of those acclimated at 15°C were then moved to 0°C for 15 days. The larvae acclimated at 15°C for 15 days were in deep diapause and accumulated little glycerol, while larvae acclimated at 15°C for 60 days were nearly ready to emerge from diapause and accumulated glycerol at 155.5 μmol/g. When larvae acclimated to 15°C for 15 days were transferred to 0°C, glycerol accumulation was stimulated to the same extent (ca 140 μmol/g) as it was in larvae that were acclimated to 15°C for 60 days and then transferred to 0°C. These results indicate that low temperature has a cumulative effect on glycerol production in larvae at different phases of diapause development. Glycerol accumulation was accomplished by activation of glycogen phosphorylase and inhibition of fructose-1,6-bisphosphatase, and activation of enzymes associated with glycerol synthesis, mainly glyceraldehyde-3-phosphatase and polyol dehydrogenase with glyceraldehyde activity.     

Metabolic dynamics across prolonged diapause development in larvae of the sawfly,Cephalcia chuxiongica (Hymenoptera: Pamphiliidae)

Nutrient metabolism is crucial for the survival of insects through the diapause. However, little is known about the metabolic mechanism of prolonged diapause. The sawfly, Cephalcia chuxiongica (Hymenoptera: Pamphiliidae), is a notorious defoliator of pine trees in southwest China. One of the distinguishing biological characteristics of this pest is the prolonged diapause of about 1.5 years. In this study, the body lipid, carbohydrate (total body sugar, glycogen, trehalose, and glucose), protein, and glycerol contents were measured in diapausing larvae of C. chuxiongica. The results showed that the changes of biochemical composition in C. chuxiongica are associated with the diapause initiation, maintenance, and termination phases. During the initiation phase, trehalose, glucose, and glycerol increased significantly, but glycogen decreased sharply. In general, the lipid, carbohydrate, and glycerol levels decreased gradually across the maintenance phase. At termination phase, the contents of glycogen and lipid persistently decreased, while an increase of trehalose, glucose, and glycerol contents were detected. The protein level was significantly higher at maintenance phase than at initiation and termination phases. It was also found that elevation of trehalose, glucose, and glycerol contents occurred in winter. These implies that the metabolites with altered levels in diapausing larvae of C. chuxiongica are responsible for maintaining a prolonged development and overwintering.     

THE SIGNIFICANCE OF THE LIGHT AND DARK PHASES IN THE PHOTOPERIODIC CONTROL OF DIAPAUSE IN METATETRANYCHUS ULMI KOCH

The daily cycle of illumination is one of several agencies which control the onset of diapause in Metatetranychus ulmi. Both light and dark phases in the cycle are concerned in the determination process.
In general, a long light phase tends to suppress and a long dark phase to induce a diapause. In any combination, the path of development is decided by the balance between diapause-preventing (light phase) and diapause-inducing (dark phase) stimuli. However, as their effectiveness does not increase linearly with duration, the existing balance changes with the phase duration.
The effectiveness of the light phase in suppressing diapause increases most rapidly between 8 and 16 hr.; that of the dark phase rises very sharply between 8 and 12 hr. Longer dark periods of up to several days duration also induce diapause but are no more effective than a 13 hr. phase. The inclusion in the cycle of very long periods of light or darkness may also influence diapause by reducing the number of complementary phases experienced by the mite during the sensitive period of development.
M. ulmi is highly insensitive to the interruption of effective light and dark phases by short intervals of darkness or light—a further indication of the slow inception of the light-and dark-phase reactions.
These findings are discussed in terms of hypothetical mechanism involving cumulative synthesis and removal of some active substance, but the experimental results cannot yet be fully reconciled with a simple hypothesis of this kind.     

Biphasic effect of low temperature on completion of winter diapause in the onion maggot,Delia antiqua

The effect of low temperature on completion of winter diapause was investigated in the onion maggot, Delia antiqua (Diptera: Anthomyiidae). Diapause was completed under constant diapause-inducing conditions of 15 degrees C and 12L-12D, without any exposure to lower temperature. The pupal period for 50% adult emergence was 117 days. None of the cold treatments at 5.6 degrees C examined in the present study advanced adult emergence; on the contrary, they delayed it. Detailed analyses of the results revealed that diapause development in D. antiqua comprises two phases which differ in sensitivity to low temperature, with the phase shift occurring at around day 60 at 15 degrees C and 12L-12D. In the first phase of diapause development, low temperature (5.6 degrees C) had no effect on diapause development. In the latter phase, by contrast, diapause development was retarded in proportion to the duration of cold treatment.     

Possible involvement of distinct photoreceptors in the photoperiodic induction of diapause in the flesh fly Sarcophaga similis

Physiological characteristics of the photoreceptors involved in the photoperiodic induction of diapause were investigated in the flesh fly Sarcophaga similis. Both the early and late phases of scotophase were sensitive to light and a light pulse during each of these phases prevented diapause. Certain physiological differences between the phases were, nevertheless, detected. Compared with early scotophase, late scotophase required a light pulse with a long period and a large number of night interruption photoperiodic cycles in order to effectively prevent diapause. The diapause-averting effects of a light pulse during early scotophase were canceled by an additional long dark period, but those during late scotophase were not. Thus, the diapause-averting effects produced during early scotophase are different to those produced during late scotophase. The early scotophase was sensitive to light at wavelengths of 470 nm or shorter, but not to light of 583 nm or longer. In contrast, the late scotophase was sensitive to light of a broad range of wavelengths, ranging from 395 to 660 nm. Furthermore, the early scotophase was considerably more sensitive to monochromatic light with low photon flux density than the late scotophase. These results suggest that different types of photoreceptor are involved in the photoperiodic response.     

Oxygen consumption in relation to sorbitol utilization at the termination of diapause in eggs of the silkworm,Bombyx mori

Rates of oxygen consumption were followed throughout the entire period of diapause in eggs of Bombyx mori. In non-diapause eggs at 25 degrees C, O(2) uptake was divisible into three phases, corresponding to morphogenetic processes. In diapause eggs at 25 degrees C, O(2) uptake showed a peak (100 &mgr;l/g eggs/h) at 1 day and then suddenly dropped to reach a level of 8-10 &mgr;l/g eggs/h at 10 days and thereafter. To break diapause, eggs were exposed to 5 degrees C for varying periods. When O(2) uptake was measured at 5 degrees C, it remained at 6 &mgr;l/g eggs/h. When eggs were chilled for increasing periods and O(2) uptake was measured immediately after warming to 25 degrees C, the rates increased after a lag phase. In HCl-treated eggs, O(2) uptake increased immediately after acid-treatment. In all cases, highly increasing O(2) uptake at 25 degrees C coincided with termination of diapause. These results were discussed in relation to sorbitol utilization at the termination of diapause.     

Effects of extending the light phase on diapause induction in a Japanese population of the two-spotted spider mite, <Emphasis Type="Italic">Tetranychus urticae</Emphasis>

Artificial lighting is a merit of a ‘plant factory’, which might be utilized to suppress an increase in pest population. We investigated the effects of extending the light phase on diapause induction in the two-spotted spider mite (TSSM), Tetranychus urticae. TSSM were reared at 18°C under light phases ranging from 2 to 64 h combined with a constant dark phase of 16 h in aluminum bottles, with white light emitting diodes attached inside to minimize fluctuations in air temperature between the light and dark phases. Diapause was induced in adult TSSM females when the light phase was 24 h or shorter, and diapause induction was inhibited when the light phase extended over 32 h. The development of deutonymphs was delayed under a diapause-inducing photoperiod. Diapause inducing photoperiods may suppress an increase in the TSSM population, by slowing down development and reproduction.     

The hormonal regulation of the larval diapause in the codling moth, Laspeyresia pomonella (Lep. Tortricidae)

The hormonal control of the facultative diapause of the codling moth has been investigated. The diapause can be divided into 4 phases or periods: (1) diapause induction by short-day conditions (SD) in young larvae, (2) initiation of the diapause in the early last larval instar by a high titre of juvenile hormone, (3) onset and maintenance of diapause with inactivity of the neuroendocrine system, as evidenced by the results of neck-ligation experiments, (4)termination of diapause by the production of ecdysteroid.Diapause-induced larvae pupated after spinning the cocoon, if the state of induction was changed by injection with the anti-juvenile hormone precocene II at the beginning of the last larval instar and subsequent results of neck-ligation experiments, (4) termination of diapause by the production of ecdysteroid. treated with juvenile hormone during the first 1.5 days after the last larval moult and subsequently reared under SD. Under LD, continuous application of juvenile hormone during the last larval instar and after spinning did not prevent the insects from moulting to either a supernumerary larva, a pupa or a larval-pupal intermediate. Termination of diapause, i.e. pupation, was achieved by injecting diapausing larvae with 20-hydroxyecdysone. Although juvenile hormone was found to have a prothoractropic effect in diapausing larvae, no pupal moult could be induced by the application of the hormone. Contrary to the hormonal situation before pupation of nondiapausing larvae, no juvenile hormone could be detected before or during the pupation of larvae after diapause.     

Diapause in a Glasgow strain of the flour moth, Ephestia kuehniella

ABSTRACT. The incidence of diapause in Ephestia kuehniella Zeller from an unhealed granary in Scotland was influenced by both photoperiod and temperature. At 25°C, nearly 50% of larvae entered diapause when reared in continuous darkness (DD) and up to 30% did so in short photoperiods. Little diapause was detected around LD 14:10 but a second, smaller peak of about 20% occurred at LD 16:8 and LD 18:6, falling away again to nearly zero in continuous light. More larvae entered diapause when reared continuously at 15 or 20°C than at 25 and 30°C. However, when larvae reared from hatch at 25°C in LD 16:8 were transferred after 1 week to 15°C in LD 9:15, almost twice as many entered diapause as did those reared at 15°C throughout. The sensitive phase for diapause induction occurred near the start of the final instar. The mean duration of diapause was between 2 and 3 months in most photoperiods at 20 and 25°C, and was shorter at 15°C. However, in DD at 25°C, it lasted about 7 months. Termination of diapause was hastened in larvae reared at 25°C in DD by transferring them to LD 14:10, and also by chilling them at 7.5°C for 6 weeks before returning to 25°C in DD. In an unhealed store in southern England, viable adults emerged from May to July and originated from larvae which terminated diapause relatively late. It would appear from the results of transferring larvae back to the laboratory at various times during the winter that some phases of diapause development were completed quite early after exposure to low temperatures, although no further development took place in the store until temperatures rose again in April.     

Adult diapause in the Colorado potato beetle, Leptinotarsa decemlineata: effects of external factors on maintenance, termination and post-diapause development

ABSTRACT. This study tests the effects of temperature, water, food and photoperiod on breaking diapause and resumption of activity (emergence from the soil) in Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae) diapausing under natural winter conditions or at constant temperatures in the laboratory. Behavioural and physiological criteria were used to characterize the various phases of diapause. Three successive phases are distinguished during hibernation: 'diapause development' or true diapause, a facultative 'post-diapause quiescence' and, finally, a transient phase of post-diapause development leading to emergence from the soil. Diapause development is completed within 3 months in the field and its duration depends on temperature. Although they are buried in the soil during this phase, beetles remain sensitive to photoperiod when artificially exposed to it. They do not emerge from the soil when exposed to higher temperatures. Thereafter, they stay in a quiescent state maintained by low temperature, low humidity or lack of food. The response to temperature changes during hibernation. In the soil, activity begins when soil temperature reaches 4–5C, but this temperature is too low to permit postdiapause development. The transient phase has a temperature threshold between 8 and 10C, whereas emergence from the soil occurs only when the temperature exceeds 11C. Post-diapause development is influenced strongly by temperature and humidity. After emergence, post-diapause development leads eventually to reproduction. Food is essential for reproduction after diapause whereas photoperiod plays no further role.     

Multiple role of temperature during insect diapause: a review

The review emphasizes that there are multiple pathways to diapause completion. The programmed course of events is modified by environmental cues. Often chilling is not a prerequisite for the completion of hibernation diapause (examples tabulated). Diapause completion progresses well at intermediate or high temperatures, sometimes it is even stimulated by high or increasing temperature. Low temperatures are important, as they (1) conserve metabolic reserves, (2) prevent resumption of post-diapause morphogenesis and thus synchronize the life-cycle, (3) represent contrast to the later increase in temperature. Diapause consists of phases with different prerequisites. There is a principal difference between diapause development and photoperiodic activation as indicated by the subsequent physiological condition of insects.