Seasonal variation in embryonic diapause of the striped ground cricket, Allonemobius fasciatus

ABSTRACT. The incidence and intensity of embryonic diapause in the striped ground cricket, Allonemobius fasciatus DeGeer (Gryllidae), is influenced by the date of oviposition and the incubation temperature. At 20C, embryos develop normally until the end of the appendage-formation stage when diapause occurs. The intensity of 'winter' diapause at this stage is greatest in eggs laid early in the season. Incubation at higher temperature induces a different and earlier 'summer' diapause. The higher the temperature the earlier the stage at which this developmental suppression is imposed. At 27 and 30C, some individuals develop without any interruption and hatch within 20 days. The proportion of these fast-developing eggs is low in summer but increases towards the end of the laying season. In nature, this species is univoltine and precocious hatching before winter is unlikely.
When embryos diapausing at an early stage at 27C are transferred to low temperatures, they resume development and enter winter diapause. The time required to reach the winter-diapause stage after transfer is inversely related to the temperature (30-30C). At 30C, sunmer diapause is neither induced nor maintained.     

Oviposition date and pattern of embryogenesis in the grasshopper Chorthippus brunneus (Orthoptera, Acrididae)

The morphological development of eggs of the grasshopper Chorthippus brunneus , laid during successive periods through a single season, was closely monitored in the field at a site in NW England. Eggs within the same clutch developed synchronously. However, there were significant differences between clutches laid at the same time, reflecting spatial variation in microclimate (and possibly maternal effects and/or genetic differences). Eggs laid early in the season entered diapause and hatched relatively synchronously in the following spring. Eggs laid later failed to reach the diapause-stage before winter, but hatched, albeit late and with greater variance. Diapause was averted in these eggs. Plasticity in the nature of diapause has been documented from laboratory studies of many insect species (including C. brunneus ,) but this study directly relates the incidence of such developmental plasticity to oviposition date and seasonal temperature cycles in the field. The adaptive significance of diapause aversion and the evolutionary consequences for female reproductive behaviour are considered.     

Influence of temperature on hatching of winter eggs of fruit-tree red spider mite, Panonychus ulmi (Koch)

The effects of the duration and degree of chilling, and the temperature of incubation, on hatching of winter eggs of Panonychus ulmi (Koch) were investigated. For chilling, 0°C and 5°C were more effective than — 5° and 9°, and the limits for the reaction were close to — 10° and 15°. As the chilling period was increased from 60 to 200 days, the percentage hatch on incubation at 21° increased, and the mean incubation time and its variance decreased. Before the maximum effect of chilling was achieved, percentage hatch on incubation at 9° and 15° was higher than at 21°; 27° was lethal to most winter eggs though not to summer eggs. After chilling, the later stages of diapause development could occur at temperatures from 0° to 21°₎ i.e. above and below the threshold temperature for morphogenesis, 6–7° in both winter and summer eggs. Diapause development cannot, therefore, be a unitary process. The significance of the results is discussed in relation to forecasting the time of hatch in the field, and to the phenological aspects of hatching in the spring.     

Diapause development in frozen larvae of the goldenrod gall fly, Eurosta solidaginis fitch (diptera: tephritidae)

Seasonal changes in metabolic rate and the potential for morphological development demonstrated that third-instar larvae of the goldenrod gall fly, Eurosta solidaginis Fitch, exhibit a distinct winter diapause. Metabolic rate (CO2 production) was significantly lower from 15 October to 9 February than in early autumn (9 September) and spring (1 March) samples. The induction of diapause coincided with the development of cold-hardening, maximum larval mass, and gall senescence, but our experiments did not identify specific cues triggering diapause induction. We examined the influence of exposure to 0 degrees C and -20 degrees C on diapause development. Diapause development in larvae stored at 0 degrees C occurred at approximately the same rate as in nature. Until 15 December the larvae were in the refractory phase of diapause (incapable of morphological development, even at permissive temperatures), but afterward moved to the activated phase within which diapause intensity decreased until termination in February. Diapause development occurred in larvae collected during the winter and stored at -20 degrees C for periods of 1 week to 3 months. Diapause intensity decreased in frozen larvae through the winter but at a slower rate than in larvae stored at 0 degrees C.     

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 temperature on the termination of prolonged larval diapause in the chestnut weevil Curculio sikkimensis (Coleoptera: Curculionidae)

Adults of the chestnut weevil Curculio sikkimensis emerged over a 3-year period under laboratory and quasi-field conditions due to a prolonged diapause that occurred at the mature larval stage. Variable proportions of the larvae remained in diapause after a single cold (5 degrees C) treatment of 120 days. Extension of the chilling period to as long as 540 days did not increase the percentage of diapause termination, and excessively long chilling actually reduced the percentage. Chilling was not indispensable to the termination of larval diapause. Diapause intensity was very high and variable, and more than 1000 days at 20 degrees C was necessary to reactivate all diapause larvae. When the diapause larvae were exposed to cycles of low (5 degrees C for 120 days) and high (20 degrees C for 240 days) temperatures, the percentage of diapause termination reached 100% after two or three such cycles. Thus, the prolonged diapause of C. sikkimensis has characteristics similar to the common short winter diapause in other insects, but has unique characteristics that ensure polymodal reactivation over several years.     

Life history strategy and egg diapause in the intertidal collembolan Anurida maritima

ABSTRACT.

• 1 In a field survey of 1 year the intertidal collembolan Anurida maritima (Guérin) was found to be a univoltine species. Every year in early May a new generation emerges from eggs that overwinter. The first animals become adult in July. Almost all eggs laid in summer do not hatch, but undergo a period of diapause. Diapause is terminated in autumn by temperatures below 5°C. However, due to the low temperatures in winter further egg development is suppressed until spring.
• 2 The adult animals die, mainly in autumn. One of the causes of mortality may be starvation. In late autumn body size decreases and glycogen and lipid content are lower.
• 3 It is argued that in autumn due to low temperatures A.maritima, which forages during low tide on the open shore and seeks refuge underground before the incoming tide, is too sluggish to find food in the limited period of low water. This probably explains why this cosmopolitan species has developed a strategy to survive the winter in the temperate zone in the egg stage.

     

Thermal response and reversibility of prolonged larval diapause in the chestnut weevil, Curculio sikkimensis

Curculio sikkimensis undergoes prolonged larval diapause that is terminated by chilling and warming cycles. To examine the effects of warming temperatures and their duration on diapause termination, we exposed diapause larvae that had not been reactivated after chilling at 5 °C to 20 or 25 °C and chilled them again before incubation at 20 °C. With increasing warming duration at 20 °C, diapause termination after chilling increased and shorter chilling durations became effective. In contrast, few or no larvae warmed at 25 °C terminated diapause after chilling, irrespective of the warming duration. To investigate the effect of warming temperature on diapause intensity, larvae with diapause weakened by initial incubation at 20 °C after the first chilling were subsequently incubated at 15, 20, or 25 °C, then chilled at 5 °C before incubation at 20 °C. Diapause termination increased significantly after the larvae were treated at 15 or 20 °C but decreased significantly after they were treated at 25 °C. The intensification of prolonged diapause at 25 °C was reversed when the larvae were transferred to 20 °C. Diapause intensity in C. sikkimensis therefore decreases at 20 °C, increases at 25 °C, and can be reversed by alternately exposing diapause larvae to 20 and 25 °C. In C. sikkimensis, prolonged diapause does not always proceed in one direction, and its intensity fluctuates in response to ambient temperature conditions.     

Developmental arrest during embryonic development of the common chameleon (Chamaeleo chamaeleon) in Spain

Embryonic development of the common chameleon, Chamaeleo chamaeleon, was monitored from oviposition to hatching at a field site in southwestern Spain and in the laboratory under five experimental temperature regimes. Embryos were diapausing gastrulae at the time of oviposition; developmental arrest in the field continued as cold torpor during winter. Postarrest development in the field commenced in April, and hatching occurred in August, for a total incubation period of 10.5 mo. In the laboratory, one group of eggs was incubated at a constant warm (26 degrees C) temperature. The remaining treatments simulated field conditions and consisted of initial periods of warm temperature of 0, 27, 46, and 71 d, a subsequent 4-mo period of cold winter (16 degrees C) temperature, and a final period of warm (26 degrees C) temperature. Embryos in the constant warm temperature treatment were in diapause an average of 3 mo, with clutch means ranging from 2 to 4 mo. Hatching among clutches occurred over 2 mo. In contrast, for field and experimental eggs that experienced cold winter conditions, hatching within treatments occurred over 2-14 d; "winter" conditions synchronized development. The length of time between the end of cold conditions and hatching did not differ among treatments; development thus resumed as soon as temperature was suitable regardless of the initial period of warm temperature. Diapause in nature thus insures that embryos remain gastrulae after oviposition despite nest temperatures that may be warm enough to support development.     

Effect of temperature on embryonic development and diapause in Chesias legatella (Lepidoptera: Geometridae)

The changing temperature requirements of Chesias legatella Schiffermüller during embryonic development are investigated. The eggs are laid in the autumn and embryogenesis is interrupted by a short diapause at the germ band stage, after which development proceeds slowly throughout the winter. The effects of temperature during the pre-diapause, diapause and post-diapause phases of development are assessed by examination of the rate and variability of morphogenesis through all the embryonic stages. An examination is presented of the effect of temperature during the late autumn on the rate and uniformity of diapause termination, and consequently on the length of the hatching period the following spring.     

Evidence for reversible change in intensity of prolonged diapause in the chestnut weevil Curculio sikkimensis

The chestnut weevil Curculio sikkimensis undergoes a prolonged larval diapause that is completed by repeated exposure to chilling and warming. We examined the possible reversibility of diapause intensity in response to temperature changes. All larvae were subjected to an initial chilling followed by incubation at 20°C to force pupation of the 1-year-type larvae that require only one winter for diapause completion. We then exposed the larvae remaining in prolonged diapause to a second chilling at 5°C for different lengths of time, preceded or not preceded by incubation at 20°C (moderately high) and/or 25°C (high) and followed by a final post-chilling reincubation at 20°C. Many of the prolonged-diapausing larvae subjected only to a brief second chilling were re-activated upon reincubation. However, short exposure to 25°C before this second chilling dramatically decreased the percentage of larvae completing diapause. When larvae were exposed to 25°C for a short period, then incubated at 20°C and subjected to the brief second chilling, many were re-activated during reincubation. The chilling time required for most of the larvae to complete diapause decreased after pre-chilling incubation at 20°C and increased after incubation at 25°C. These results demonstrate that diapause intensity in C. sikkimensis changes reversibly in response to changes in ambient temperature.     

Repeated cycles of chilling and warming effectively terminate prolonged larval diapause in the chestnut weevil, Curculio sikkimensis

Curculio sikkimensis (Coleoptera: Curculionidae) requires one or more years to complete its life cycle, owing to prolonged larval diapause. To compare the effects of temperature cycles and total periods of chilling on the termination of prolonged diapause, larvae were subjected to different chilling (5 degrees C) and warming (20 degrees C) cycles ranging from 30 to 720 days, and all cycles were repeated until the sum of chilling and warming periods reached 720 days. The prolonged diapause of C. sikkimensis was more effectively terminated by repeated cycles of chilling and warming than by prolonging the continuous chilling period. However, extremely short temperature cycles were not highly effective in enhancing diapause termination, even when such cycles were repeated many times. To examine the role of warming periods on diapause termination, diapause larvae were subjected to a sequence of chilling (120 days at 5 degrees C) and warming (240 days at 20 degrees C) with a warming period (0-120 days at 20 degrees C) inserted in the chilling period. Diapause larvae that were not reactivated in the first chilling period required exposure to a certain period of warming before they were able to complete diapause development in the subsequent chilling. Thus, C. sikkimensis appears to spread its reactivation times over several years in response to seasonal temperature cycles.     

Reversible change in embryonic diapause intensity by mild temperature in the Chinese rice grasshopper, Oxya chinensis

The effects of temperature on maintenance and termination of embryonic diapause were investigated in Jining (35.4°N, 116.6°E) and Sihong (33.5°N, 118.2°E) strains of the Chinese rice grasshopper, Oxya chinensis Thunberg (Orthoptera: Catantopidae). Eggs of both strains entered diapause when incubated at 30, 25, or 20 °C. Chilling at 8 °C had an evident effect on diapause termination and almost all eggs chilled for 60 days ended diapause development. Chilling of eggs at 8 °C for only 20 days failed to result in any hatching at 20 °C, suggesting that such level of chilling was not enough to induce diapause termination. However, the treatment combining incubation of eggs at 30 °C for varying lengths of time with subsequent incubation to 20 °C had a distinct effect on the completion of diapause of the eggs. The results indicate that there were two temperature optima, that is, low temperature (chilling) and high temperature, for diapause development in this grasshopper species. Incubation of chilled eggs at 20 °C for 5–15 days followed by further incubation at 25 °C reduced termination of diapause significantly compared with the eggs only chilled at 8 °C. Exposure of eggs chilled at 8 °C to a pulse of 25 °C from 1 to 7 days, separated by a 20-day interval at 8 °C, resulted in a decrease in the percentage of successfully hatched eggs as the length of the pulse of 25 °C increased. The results suggest that diapause intensity may be restored at moderately high temperatures. This reversible change in diapause intensity would play an important role in maintaining diapause before winter.     

Egg development and diapause: ecophysiological and genetic basis of phenological polymorphism and adaptation to varied hosts in the green oak tortrix,Tortrix viridana L. (Lepidoptera: Tortricidae)

Experiments concerned 8 tortrix populations associated with varied oak species. They showed that the egg development included a phase of diapause. Completion of embryogenesis at 20 degrees C was used as a criterion for whether diapause was completed. Under semi-natural conditions diapause terminated in late autumn or early winter, then eggs developed continuously, without postdiapause winter quiescence, even in severe cold. The eggs from the populations associated with holm or cork oak completed diapause then hatched later than those from the populations associated with sessile or pubescent oak. Reciprocal crossbreedings confirmed that this phenological polymorphism was genetically determined. Under constant temperatures the physiological state of diapausing eggs, assessed by measuring their cold requirements to complete diapause by exposure to 8 degrees C, varied gradually with increasing age. This diapause development was strongly temperature-dependent. Cold requirements of diapausing eggs were much higher in a late-hatching than in an early-hatching population. This explains the phenological polymorphism of the tortrix: the more eggs need cold, the later they complete diapause in autumn, and the later they hatch in spring. Egg cold requirements varied widely within populations too, which resulted in large variations in the date of diapause end among individuals. The date of egg hatch was influenced by the temperatures occurring during diapause and postdiapause, but apparently not by photoperiod.     

Effect of anoxia on diapause termination in eggs of the false melon beetle, Atrachya menetriesi

The effect of anoxia on diapause development in the leaf beetle Atrachya menetriesi was investigated to elucidate the role of oxygen in regulation of egg diapause. While anoxia alone had no effect on diapause termination, it decreased diapause intensity before chilling. Such an effect reached a maximum level when anoxia lasted for about 10 days. Anoxia applied during the pre-diapause stage also reduced diapause intensity. On the other hand, anoxia terminated diapause when the diapause intensity had been lowered by sufficient duration of chilling (50 days at 7.5 degrees C). The effect of anoxia was temperature dependent; the larger effect was elicited when anoxia was combined with a higher temperature. A 50-day chilling caused more than 20% of eggs to terminate diapause upon transfer to warm conditions. However, when this chilling period was interrupted on the 20th day by a 5-day exposure to a high temperature of 20, 25 or 30 degrees C, the effect of the former chilling was cancelled partially or completely, suggesting that warming reversed diapause development. This reversing effect of a high temperature, however, was not manifested when the warming was combined with anoxia. The results suggest that anoxia inhibits diapause reversal and facilitates a certain process of diapause development. The sequence of exposure to anoxia and chilling is not important.     

Demarcation of diapause development by cold and its relation to time-interval activation of TIME-ATPase in eggs of the silkworm, Bombyx mori

We investigated the mode of action of winter cold in the termination of diapause by investigating Time-Interval-Measuring Enzyme (TIME). First, we determined the period of cold required for the completion of diapause development. Synchronously developing egg batches of a pure strain (C108 Bombyx mori silkworm) were used to minimize variations in hatching time. Hatching occurred with only 18 days chilling at 5 degrees C when the incubation at 25 degrees C after the chilling was elongated. The 18-day period was much shorter than we expected; diapause in B. mori is known to terminate completely with about 100 days of chilling. Even in such a short period of chilling, no sporadic hatching occurred. Moreover, we determined that a temperature-insensitive stage, which we called "Neboke", followed the short cold-requiring stage. Thus, the stage of diapause development was demarcated from other stages of diapause. While the length of diapause development was elongated when chilling was delayed after oviposition, the Neboke stage length was invariant. Cold evidently exerts its effect only on diapause development. When TIME was purified from eggs and chilled in test tubes, a transitory burst of its ATPase activity occurred at a time equivalent to shortly before the completion of diapause development; this was an interval-timer activation. The mechanism by which cold activates TIME to measure the time interval may help explain in biochemical terms the insect's adaptation to its seasonal environments.     

中华稻蝗长沙种群的生活史及其卵滞育的进化意义

中华稻蝗Oxya chinensis为重要的水稻害虫,在我国除青海、西藏、新疆、内蒙古等未见报道外,南起海南北至东北均有分布,在许多的分布区域1年发生1代。为探索中华稻蝗长沙种群的生活史及其季节适应特征,通过野外和实验室的研究,调查了其发生代数、孵化率和卵滞育率的季节变化及越冬卵的存活率。结果显示,中华稻蝗长沙种群为1年2代和1年1代混合发生:第1代卵产卵后大部分孵化为若虫而1年完成2代,但亦有19.4% -4.1%的卵不孵化而1年只能完成1世代。第1代成虫于6月上旬至8月上旬羽化,6月下旬至8月中旬产卵;第2代若虫于7月初开始孵化,9-10月羽化为成虫,10月上旬至11月下旬产卵。在室外自然条件下,中华稻蝗长沙种群6-8月(第1代)和10-11月(第2代)所产卵块均为部分滞育,滞育率为30%左右,皆无显著差异。然而,其卵滞育率在12月以后显著降低,仅为6.6%或以下,卵滞育快速地得以解除。因此,包括非滞育卵和滞育解除卵,中华稻蝗长沙种群的越冬卵皆以非滞育状态度过其后的寒冷季节。即使是遭遇长江流域2007年末至 2008年初异常寒冷的冬季,在长沙地区越冬后其卵的存活率亦在98%以上。非滞育状态的中华稻蝗长沙种群越冬卵完全能安全地越冬,其滞育的发生并非是为了提高其耐寒性而安全度过不适环境。并探讨了中华稻蝗长沙种群卵滞育的进化意义。     

Geographical variation in Japan in egg development of the mayfly, Ephoron shigae (Ephemeroptera: Polymitarcyidae)

1. The effect of temperature on embryonic development was compared in four populations, two bisexual and two unisexual, of Ephoron shigae , including one each near the northern and southern periphery of the species range in Japan.
2. Eggs from every population were chilled at 4, 8 or 12 °C for diapause development after 50 days at 20 °C for pre-diapause development (experiment I). Some eggs hatched during chilling at 8 °C or 12 °C, whereas no eggs hatched at 4 °C. The rate of hatching in a given condition of chilling was higher for the eggs from warmer winter environments.
3. Chilling at 4 or 8 °C effectively facilitated diapause development. Chilling at 12 °C was, in general, not so effective, but relatively effective for the eggs from warmer winter environments.
4. Eggs were incubated at 8, 12, 15 or 20 °C after chilling at 4 °C to examine the effect of temperature on post-diapause development (experiment II). The eggs incubated at higher temperature after chilling hatched quicker and more synchronously and had higher hatching success.
5. The relationship between temperature and the days required for hatching after chilling was well described by the power function. There was no significant difference in the slope of the regression lines (i.e. temperature dependency) among local populations. However, a longer time was required for hatching at a given temperature for the population from the colder winter environment.
6. There was no detectable difference in the observed intraspecific variations between unisexual and bisexual populations.     

Embryonic stage of obligatory diapause and effects of abiotic conditions on egg hatching in the balsam twig aphid,Mindarus abietinus

Diapause‐mediated dormancy in overwintering insect eggs has rarely been studied with regard to the ecological factors controlling postdiapause development. In insects of temperate latitudes, water availability at the end of winter, in interaction with temperature, could control the resumption of development for insect stages in postdiapause quiescence. The balsam twig aphid, Mindarus abietinus Koch (Hemiptera: Aphididae), overwinters as eggs in southern Québec, Canada, on balsam fir, Abies balsamea (L.) Miller (Pinaceae), in Christmas tree plantations, where it is known as a pest. Previous work has shown that eggs of this aphid maintain low water content during winter, presumably to survive sub‐zero temperatures. Conversely, in late winter and early spring, they passively or actively absorb surrounding moisture, which is accompanied by notable changes in size, shape, and fresh mass. The primary objective here was to determine the embryonic stage at which winter diapause starts and is maintained in M. abietinus, a relatively primitive aphid. Secondly, we tested the hypothesis that free water availability to postdiapause eggs, in combination with temperatures above developmental threshold, is essential for embryonic development and hatching, by experimentally soaking field‐collected eggs in water at controlled frequencies. We observed that embryogenesis starts at the time of egg laying and stops after a few days, before the anatrepsis stage of blastokinesis is complete, when the germ band has not yet entirely immersed itself into the yolk. We also found that water surrounding overwintered eggs on fir shoots, in interaction with temperature regime, significantly increases M. abietinus egg hatching rates. Potential impacts of environmental factors such as precipitation are discussed in relation to M. abietinus egg hatching rates and potential for population growth in spring.     

The effects of temperature and moisture on the inception of diapause in eggs of the Australian plague locust,Chortoicetes terminifera Walker (Orthoptera: Acrididae)

Chortoicetes terminifera lays a mixture of diapause and non-diapause eggs during the autumn months of March and April. Diapause eggs cease development in late anatrepsis when kept in moist soil at 32.5°C. These conditions favour rapid embryogenesis and hatching in non-diapause eggs. Significant reductions in the incidence of diapause were obtained when newly-laid eggs were kept for 40 days at either 15.5°C or 32.5°C in conditions that prevented the uptake of water. In eggs held for a similar period in moist soil at 15.5°C, diapause was virtually eliminated. These eggs did not develop to the stage at which water uptake occurs. It is suggested that the inception of diapause is dependent upon the occurrence of warm, moist conditions at the time of laying and that these requirements are an integral part of the overall inductive process, which also involves the temperature and photoperiod at which the parent insects are reared. A flow-diagram illustrating the development of diapause eggs in relation to temperature and moisture is discussed in relation to the survival of the species.