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.     

Bimodal life cycle of the burying beetle Nicrophorus quadripunctatus in relation to its summer reproductive diapause

Abstract 1. Under natural conditions in Kyoto, Japan, the reproductive activities of Nicrophorus quadripunctatus Kraatz (Coleoptera: Silphidae) decreased in summer and the species showed a bimodal life cycle.
2. In the laboratory, most adult pairs raised at 20 °C under a LD 12:12 h regime reproduced when provided with a piece of chicken. In adults raised at 20 °C under a LD 16:8 h regime, however, both reproductive behaviour and ovarian development were reduced. It is concluded that these adults entered a reproductive summer diapause.
3. High temperature (25 °C) also suppressed the reproductive behaviour even under a favourable LD 12:12 h regime. In the field, therefore, adults reduce their reproductive activity in summer because of diapause induced by long-day photoperiods and direct inhibition of reproduction by high temperatures.
4. When the temperature was changed from 20 °C to 25 °C immediately after hatching of larvae, they reached the wandering stage in 95% of adult pairs. When the temperature was changed from 20 °C to 25 °C immediately after oviposition, however, no larvae hatched in 85% of pairs. Egg mortality was significantly higher at 25 °C than at 20 and 22.5 °C; no eggs hatched at 27.5 °C. The physiological mechanisms for reducing reproduction probably prevent the beetles from inefficient oviposition in summer.     

Effects of temperature during chilling and pre-chilling periods on diapause and post-diapause development in a katydid, Eobiana engelhardti subtropica

In Eobiana engelhardti subtropica, early laid eggs reach the diapause stage in early autumn. For long periods before winter, the eggs are exposed to temperatures higher than their theoretical lower threshold for development. In contrast, late-laid eggs cannot reach their diapause stage before winter. Our study showed that E. e. subtropica copes with these difficulties via the thermal response involving embryonic diapause. In this katydid, the almost fully developed embryo undergoes an obligatory diapause. When diapause eggs were maintained at a temperature of 20 degrees C or higher, diapause persisted for a long time. Diapause was effectively terminated by temperatures ranging from 1 to 11 degrees C, and hatching occurred successfully at temperatures from 11 to 15 degrees C. In addition to the chilling temperature, pre-chilling temperature modified diapause intensity and hatching time. Diapause eggs hatched earlier after chilling when the pre-chilling temperature was lower, within a range of 14.5-25 degrees C. Thus, the low-temperature requirement for diapause termination prevents early laid eggs from untimely hatching in autumn, and low temperatures before and during winter decrease diapause intensity and shorten the hatching time in the following spring. When eggs were chilled before diapause, they tolerated chilling and averted diapause. Thus, even if eggs encounter low temperatures before diapause, they can hatch in the following spring.     

Geographic parthenogenesis in the stream mayfly Eurylophella funeralis in eastern North America

Unisexual populations were found largely at the periphery of the geographic range of the mayfly Eurylophella funeralis in eastern North America. Bisexual populations generally had a normal sex ratio but at least two populations were observed with only about 2–11% males. Hatch success for unfertilized eggs depended on temperature but in general averaged >61% in unisexual populations and < 14% in bisexual populations. Eggs took about 65 d to hatch at 10°C and 19 d at 25°C; no hatching occurred at 5 or 30°C. Successful oviposition was observed for about 97% and 20% of unmated female adults from unisexual and bisexual populations respectively. The hatch success of unfertilized eggs for three geographically distinct bisexual populations was in versely correlated with the proportion of males in the populations. The genetic structure of bisexual and unisexual populations was very similar. Unisexual populations consists of clones, with as few as four recognizable phenotypes being present at a given location. All offspring had the same phenotype as the mother. Tychoparthenogenesis is suggested as the most reasonable working hypothesis concerning the origin of unisexual populations in E. funeralis .     

Effect of temperature on the hatching time of eggs of Ephemerella ignita (Poda) (Ephemeroptera:Ephemerellidae)

SUMMARY. Eggs of Ephemerella ignita (Poda) were kept at eight constant temperatures (range 5.9–19.8°C) in the laboratory. Over 85% of the eggs hatched in the temperature range 10.0–14.2°C but the percentage decreased markedly to 39% at 5.9°C and 42% at 19.8°C. Hatching time (days after oviposition) decreased with increasing water temperature over the range 5.9–14.2°C and the relationship between the two variables was well described by a hyperbola. Therefore, the time taken for development was expressed in units of degree-days above a threshold temperature. Mean values (with 95%CL) were 552 (534–573) degree-days above 4.25°C for 10% of the eggs hatched, 862 (725–1064) degree-days above 3.57°C for 50% hatched and 1383 (1294–1486) degree-days above 3.14°C for 90% hatched. These values can be used to predict hatching times at temperatures below 14.68°C for 10% hatched, 14.54°C for 50% hatched and 14.45°C for 90% hatched. At higher temperatures, the hatching time and the number of degree-days required for development both increased with increasing temperature. Equations were developed to estimate the number of degree-days required for development at these higher temperatures.
Eggs were also placed in the Wilfin Beck, a small stony stream in the English Lake District. Maximum and minimum water temperatures were recorded in each week and the summation of degree-days was used to predict the dates on which 10%, 50% and 90% of the eggs should have hatched. There was good agreement between these estimates and the actual hatching times. Only 10–15% of the eggs hatched between October and late February with most of the eggs hatching in March, April and May. Nymphs hatching in October and November probably did not survive the winter.     

Methods for the activation of the resting eggs of Daphnia

SUMMARY. 1. The conditions required to initiate development of resting eggs of thirty-six clones of Daphnia representing seven species were investigated.
2. The temperature of both dark incubation and subsequent light treatment are shown to affect hatch success. By varying these parameters the majority of resting eggs from each test clone were stimulated to develop. Arctic clones required a low hatching temperature (7°C), whereas clones from warmer climates hatched best at 14–21°C.
3. Variation in hatching cues existed between conspecific individuals from different collection sites. These differences suggest that research determining macro- and microgeographic patterns in hatching phenology would be fruitful.     

Relationships between maternal size, egg diameter, time of spawning season, temperature, and length at hatch of Atlantic silverside, Menidia menidia

Eggs were stripped from gravid Atlantic silversides collected on two occasions, once during the early part and once during the late part of the natural spawning season. Unfertilized egg diameter was not correlated with length of the female, nor was it significantly larger during the early part of the season. Eggs were fertilized and incubated in the laboratory. Larval length at hatch was measured every 24 h during the hatching period after embryos were incubated at 18 or 25° C. Lower incubation temperature caused a significantly greater length at hatch for the offspring of each of the 20 females studies. In most cases (17 out of 20 at 25° C, 10 out of 20 at 18° C), there was a significant decrease in length at hatch during the hatching period for a given female's eggs incubated at a given temperature. In the natural environment, larvae hatched early in the season under cooler temperatures could average 12% longer than those hatched later under warmer temperatures, and therefore may have a greater chance of survival. The results help to explain the observation that field-caught M. menidia that hatched early in the season are larger at any given age than those that hatched late in the season.     

Cold-hardiness and development of eggs of Rhopalosiphum insertum

Abstract. 1. The eggs of Rhopalosiphum insertum (Walker) showed a seasonal increase in cold-hardiness under field conditions. Their supercooling point fell from -35°C in November to below -40°C in January, then rose to-35°C or above by March.
2. Laboratory experiments demonstrated that both temperature and date affected cold-hardiness of the eggs. The supercooling points of eggs kept at 16 h photoperiod or in darkness did not, however, differ significantly.
3. Eggs brought from the field into warm, long-day conditions would not hatch until after mid-January. After this date, per cent hatch was significantly greater in 16 h photoperiod than in darkness; it did not differ between eggs kept at 5 or 0°C, but was reduced at -5°C.
4. It is concluded that eggs of Rinserturn are in diapause until mid-January, and that hatching rate and cold-hardiness are determined by separate environmental factors.     

Effects of temperature of adults and eggs on the induction of embryonic diapause in the band-legged ground cricket, Dianemobius nigrofasciatus

Abstract.  Eggs laid by adult female Dianemobius nigrofasciatus , reared under long-day (LD 16 : 8 h, 25 °C) or short-day (LD 12 : 12 h, 25 °C) conditions from the nymphal stage, are kept at several constant temperatures. At 22.5–30.0 °C, eggs laid by long-day adults show lower incidences of diapause than those laid by short-day adults. In both eggs laid by adults under long-day conditions and those under short-day conditions, the higher the temperature at which the eggs are kept, the lower the incidence of diapause. When eggs of long-day adults are exposed to a low-temperature pulse (10 °C, 24 h) on the day of deposition (day 0), the incidence of diapause increases. The low-temperature pulse on day 1 does not increase the incidence of diapause. By contrast, when the eggs of short-day adults are exposed to a high-temperature pulse (35 °C, 24 h) on day 0 or day 1, the incidence of diapause decreases. The temperature pulses on day 0 are more effective at diapause prevention. Staining of diapause eggs by the Feulgen–Rossenbeck method shows that the eggs enter diapause at the blastoderm stage, which is on day 1 or day 2 at 25 °C. The exposure of adults to long days and higher temperatures prevents the eggs from entering diapause. In D. nigrofasciatus , embryonic diapause is controlled by maternal effects, adult photoperiod and temperature, and egg temperature before or at diapause.     

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.     

Hatching of cladoceran resting eggs: temperature and photoperiod

Summary 1. We identified temperature and photoperiod conditions under which the hatching of 45 cladoceran species could be elicited. Identification of appropriate hatching cues is of primary importance for the exploration of the ties between active and diapausing stages.
2. Incubation temperature affected the hatching success of resting eggs isolated from Danish, Belgian/Dutch and Spanish sediments. In general, most hatchlings and species were retrieved at 15 °C. Danish and Belgian/Dutch resting eggs hatched more successfully under a long day photoperiod than in continuous illumination.
3. Most species could be retrieved after incubation of resting eggs isolated from a limited amount of sediment (0.4 kg) under a single, well chosen combination of temperature and photoperiod. Processing additional sediment samples under seven more incubation regimes only allowed detection of 21% (Spain) to 34% (Denmark) more species.
4. The incubation period for resting eggs to hatch was strongly influenced by incubation temperature. Our results show that hatching experiments aimed at assessing cladoceran species richness and conducted at 15 °C should be continued for a period of at least 2 weeks, after which a random subset of hatchlings (e.g. n  = 100) can be selected from the total hatchling assemblage.     

Interspecific and intraspecific variations in egg hatching for British populations of the stoneflies Siphonoperla torrentium and Chloroperia tripunctata (Plecoptera: Chloroperiidae)

SUMMARY 1. The objective was to compare variations in egg hatching between the two species (interspecific variations) and between populations of the same species (intraspecific variations). There were significant interspecific, but not intraspecific, differences in female size, adult life-span, egg production, hatching success, incubation periods and hatching periods.
2. The optimum temperature for hatching success within the range 3.8–22.1°C in the laboratory and the range over which at least 50% of the eggs hatched were lower for Chloroperia tripunctata (Scopoli) (8.5°C, 4.2–17.3°C) than for Siphonoperla torrentium (Pictet) (12.8°C, 6.1–19.4°C). Few eggs hatched at 22.r°C.
3. The relationship between incubation period (d days) and water temperature (T°C) was given by: d=1219/T^1.368 for S. torrentium , d=253/T^0.459 for C. tripunctata . Both equations successfully predicted incubation periods for eggs placed in a stream. The period over which eggs hatched was much longer for C. tripunctata than for S. torrentium at all temperatures.
4. The shorter incubation period (at r>5.6°C) and shorter hatching period for S. torrentium ensure that larvae of this species are already growing when eggs of C. tripunctata start to hatch, but the prolonged hatching period of the latter species ensures a long period of larval recruitment to the population. These differences in egg hatching may reduce competition between the two closely-related species.     

Effects of temperature, sorbitol, alanine and diapause hormone on the embryonic development in Bombyx mori: in vitro tests of old hypotheses

Abstract.  An in vitro culture method is described in which embryonic development in Bombyx mori is traced at various temperatures and treatments. The results show that the induction, intensification and termination of diapause are distinct processes. Prediapause embryos, explanted from 40-h-old diapause-destined eggs and cultured in Grace's medium, continue to develop to the appendage-formation stage without arrest, which indicates that the isolated embryos have not entered diapause, whereas the development of embryos from diapausing eggs (15 days after being laid) is significantly slower. The rate of development of embryos dissected from diapause eggs increases during chilling (5 °C) and incubation (at 25 °C) gradually during chilling and dramatically at 25 °C. The in vitro experiments also reveal that sorbitol directly inhibits the development of embryos explanted from diapausing eggs but has no affect on the development of embryos from prediapause eggs. Neither alanine nor diapause hormone prevent isolated embryos from developing.     

Temperature effects on embryonic development of Paratlanticus ussuriensis (Orthoptera: Tettigoniidae) in relation to its prolonged diapause

Paratlanticus ussuriensis eggs overwinter by entering diapause, which can be prolonged to more than 1 year depending on environmental conditions. To determine temperature effects on diapause duration of P. ussuriensis eggs, the rates of embryonic development and hatching were compared at various temperatures conditions by measuring embryonic stages and egg weights. Most eggs stayed in a very young stage (blastoderm formation, stage 4) when reared at 15 and 20 °C, 10–30% eggs developed into middle or late stages when reared at 25 °C, and most embryos developed fully (stage 23/24) when reared at 30 °C. Egg weight at 30 °C was 1.5 times higher than those reared at 20 °C. Chilling induced hatching in embryos at stage 23/24. Chilling caused stage 4 embryos to develop into stage 24, but they failed to hatch in response to a second warm period. Thus, P. ussuriensis eggs can overwinter either as young embryos (initial diapause) or as fully-developed embryos (final diapause). Eggs that experience an initial diapause overwinter again the second year in a final stage diapause. The post-diapause period was shorter when embryos overwintered in a final stage diapause. The hatching rate was highest in a temperature range of 7.5–15 °C. Our results suggest that temperature is an important environmental factor for the control of prolonged diapause in P. ussuriensis and initial diapause plays an important role in the control of its life cycle.     

Assessment of temperature effects on the development and fecundity of Pullus mediterraneus (Col., Coccinellidae) and consumption of Saissetia oleae eggs (Hom., Coccoida)

Eggs, larval and nymphal periods and fecundity of Pullus mediterraneus were examined under 16 h light : 8 h dark combined with six constant temperatures: 15, 20, 25, 30, 35 and 40°C. Eggs of Saissetia oleae were used as prey. The developmental time at 15, 20, 25, 30 and 35°C was 17.23, 4.5, 2.64, 1.67, 1.28 days for eggs and 98.47, 68.88, 53.94, 28.96, 36.51 days for larval–pupal duration, respectively. At 7°C no eggs hatched, and at 40°C all the stages died after 36 h of maximum exposure except the three last stages. The fecundity of females rearing at different temperatures ranged between 1.7 eggs at 15°C and 601.86 eggs at 30°C. The pre-oviposition period ranged between 23.75 days at 15°C and 3.47 days at 35°C. The consumption of S. oleae eggs by the larvae reached 597.69 eggs during the pre-imaginal development. Females attacked more eggs than males averaging 77.69 ± 22.34 eggs per 4 day period compared with 46.97 ± 10.12 eggs per 4 day period for males.     

Embryonic development, larval growth and life cycle of Coenagrion puella (Odonata: Zygoptera) from an Austrian pond

SUMMARY. 1. Newly-laid eggs of Coenagrion puella (L.) from a pond near Herzogenburg (Lower Austria) were kept at constant water temperatures (range c .3.5°C to c .28°C)in the laboratory. Hatching success varied with temperature; no eggs hatched below 12°C and nearly all hatched at c .l6°C. Hatching time decreased with increasing temperature and the relationship between the two variables within the range 12–28 °C was well described by a power law. The length of the hatching period was less than 12 days. Hatching times estimated from the power-law equations and those obtained in the field experiments were similar. Therefore both the hatching time and the length of the hatching period in the field could be estimated from the laboratory data for the range 12–28°C.
2. The maximum number of instars from egg to imago was 11; the average body length increment (mm) per moult was proportionately constant at c .26% and Dyar's rule was applicable. The interval between moults decreased with increasing temperature up to the seventh instar and the relationship between the two variables within the range 12–28°C was well described by a power law. The moulting interval for instars 8–11 ranged from 23 to 48 days and was relatively independent of temperature. No moulting occurred at temperatures below 12°C.
3. Larval growth was logistic in the laboratory and variations in mean logistic growth rate (range 0–2.5% length day⁻¹) were related to mean temperature with no growth at temperatures <12°C. Larval growth rates in pond experiments were similar to those estimated from laboratory data, and therefore the regression equations obtained from the laboratory experiments are probably applicable to larval growth in the field.
4. Information on the life cycle of C. puella is briefly reviewed and it is concluded that C. puella from the pond near Herzogenburg has an univoltine life cycle.     

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.     

Thermal dependence of embryonic growth and development in brown trout

Fertilized eggs from a brown trout Salmo trutta population in northern Spain were incubated in the laboratory at 4, 6, 8, 10, 12, 14, 16 and 18° C. Developmental stage and embryo size were monitored by taking samples at regular intervals. Survival was maximal at 8 and 10° C and decreased at higher and lower temperatures. Despite starting development, no embryo hatched at 16 and 18° C, which suggests an upper thermal limit for development between 14 and 16° C. Time required to reach a given ontogenetic stage decreased with increasing temperature. Embryos incubated at lower temperatures were larger at 50% hatching, and these differences persisted throughout the subsequent embryonic period until the start of exogenous feeding. A comparison with previously published data indicates low interpopulation variability in thermal sensitivity of embryonic development, even in consideration of the great latitudinal range of the studies.     

Temperature dependent sorbitol utilization in diapause eggs of the silkworm,Bombyx mori

Summary A series of experiments was conducted to determine whether the interconversion of glycogen and sorbitol at the initiation and termination of diapause in eggs of the silkworm,Bombyx mori was influenced by low temperatures (5°C and 1°C). The conversion of glycogen to sorbitol and glycerol at the initiation of diapause was not affected by exposure to 5°C and 1°C. Chilling diapause eggs at 5°C for over 70 days resulted in a progressive conversion of sorbitol to glycogen. Transfer to 1°C after chilling to 5°C led to a decrease in sorbitol and glycerol conversion. While continuous chilling at 1°C completely inhibited sorbitol utilization for at least 400 days, sorbitol began to decrease immediately following transfer to 5°C. In contrast, glycerol utilization was not prevented by acclimation to 1°C, but a delayed decrease occurred. Continuous exposure to 1°C delayed hatching for a period of 440 days, whereas acclimation to 5°C resulted in 100 per cent hatching by about 100 days. A low, or high acclimation temperature 1°C or 5°C produce different effects on sorbitol utilization and termination of diapause in silkworm eggs.     

Optimal Low Temperature and Chilling Period for Both Summer and Winter Diapause Development in Pieris melete: Based on a Similar Mechanism

The cabbage butterfly, Pieris melete hibernates and aestivates as a diapausing pupa. We present evidence that the optimum of low temperature and optimal chilling periods for both summer and winter diapause development are based on a similar mechanism. Summer or winter diapausing pupae were exposed to different low temperatures of 1, 5, 10 or 15°C for different chilling periods (ranging from 30 to 120 d) or chilling treatments started at different stages of diapause, and were then transferred to 20°C, LD12.5∶11.5 to terminate diapause. Chilling temperature and duration had a significant effect on the development of aestivating and hibernating pupae. The durations of diapause for both aestivating and hibernating pupae were significantly shorter when they were exposed to low temperatures of 1, 5 or 10°C for 50 or 60 days, suggesting that the optimum chilling temperatures for diapause development were between 1 and 10°C and the required optimal chilling period was about 50–60 days. Eighty days of chilling was efficient for the completion of both summer and winter diapause. When chilling periods were ≥90 days, the durations of summer and winter diapause were significantly lengthened; however, the adult emergence was more synchronous. The adaptive significance of a similar mechanism on summer and winter diapause development is discussed.