POPULATION SEASONALITY AND LARVAL DEVELOPMENT OF LAGRIA HIRTA L. (COLEOPTERA: LAGRIIDAE)*

Abstract The seasonality of the field population of Lagria hirta L. is remarkable: adults occur only from the end of June to the beginning of August, whereas larvae exist in the whole year except June. Based on the data of field observations and larval head-width measuring, the author demonstrated that L. hirta would reproduce in a very short interval in June to August; eggs hatch and develop (molt) into the 3rd, 4th or 5th (sometimes 6th) instar before winter; after larval overwintering, the beetles complete the larval development after total 8–10 times of molts and pupate in June. The larval development rate is decreased but not completely stopped during winter (the nonparametric median test, P<0. 01). This result changes the hitherto opinion of 4 or 5-instar larval development of L. hirta and will stimulate the studies on its life history.     

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.     

Influences of daylength and temperature on the period of diapause and its ending process in dormant larvae of burnet moths (Lepidoptera,Zygaenidae)

The onset of larval diapause in the burnet moth Zygaena trifolii is clearly characterized by the larva molting into a specialized dormant morph. In a potentially bivoltine Mediterranean population (Marseille) two types of diapause can occur within 1 year: firstly, a facultative summer diapause of 3–10 weeks, and secondly, an obligate winter diapause, which can be lengthened by a period of thermal quiescence to several months in temperatures of 5°C. For the first time, three successive physiological periods have been experimentally distinguished within an insect dormancy (between onset of diapause and molting to the next non-diapause stage), using chilling periods of 30–180 days at 5°C, and varying conditions of photoperiod and temperature. These stages are: (1) a continuous Diapause-ending process (DEP); (2) thermal quiescence (Q); and finally, (3) a period of postdiapause development (PDD) before molting to the next larval instar. The result of transferring dormant larvae from chilling at 5°C to 20°C depended on the length of the chilling period. After chilling for 120–180 days, molting to the next instar occurred after 6–10 days, independent of daylength. This period corresponds with the duration of PDD. After shorter chilling periods (90, 60, 30 days and the control, 0 days) the period to eclosion increased exponentially, and included both the latter part of the previous diapause process and the 6–10 day period of PDD. However, photoperiod also influences the time to eclosion after chilling. Short daylength (8 h light / 16 h dark: LD 8/16) lengthened the diapause in comparison to long daylength (16 h light / 8 h dark: LD 16/8). Short daylength had a similar effect during chilling at 5°C, as measured by the longer time to eclosion after transfer. The shorter time to eclosion resulting from longer chilling periods (30–90 days) demonstrates that the state of diapause is continuously shortened at 5°C, and corresponds to the neuroendocrine controlled DEP. Presumably the DEP has already started after the onset of diapause. When chilling was continued after the end of the DEP, which ranged between 90 and 120 days, thermal quiescence (Q) followed (observed maximum 395 days). Different photoperiodic conditions during the pre-diapause inductive period modified diapause intensity (measured as the duration of diapause), in that a photoperiodic signal just below the critical photoperiod for diapause induction (LD 15/9) intensified diapause. Experiments simulating the summer diapause showed that PDD occurred in the range of 10–25°C. Higher temperatures (15 and 20°C) shortened the DEP at LD 16/8, so that at 20°C many individuals had already terminated diapause after 10–40 days and had molted after the 6–10 days of PDD. A temperature of 25°C unexpectedly lengthened the DEP to 110 days in several individuals. The ecological consequences and the adaptive significance of variation in the duration of the diapause are discussed in relation to the persistence of local populations predictably variable and rare climatic extremes throughout the year.     

Control of diapause in codling moth larvae

Diapause in a New Zealand strain of codling moth (Cydia pomonella Linnaeus [Lepidoptera: Olethreutidae]) was induced in larvae by photoperiods of 15 h or less. Once diapause had been initiated, it could not be terminated by any combination of conditions tested for at least 20 days after cocooning. In diapausing larvae a low rate of pupation occurred at 25 °C under a long day (18 h) photoperiod. A high rate of pupation was achieved under a long day regime when larvae were decocooned, and provided with apple as nourishment. Diapause could be terminated predictably in 94–100% of larvae by 1) conditioning at 15 °C and constant darkness for periods of 40–100 days, then 2) chilling at 2±2 °C and constant darkness for 20–50 days followed by 3) any post-chill condition periods at 25 °C, 18 h photoperiod. Complete diapause termination was achieved when 100 days conditioning was followed by 30 days or 50 days post-chill period. Under these conditions, 76% termination occurred in the post-chill period after 10 days, and 93% after 25 days.To terminate diapause in codling moth larvae, we recommend that a 100 days conditioning followed by 30 days chilling and 50 days post chilling periods be used.     

Cell cycles in embryos of the silkworm,Bombyx mori: G2-arrest at diapause stage

Summary In the silkworm, Bombyx mori, diapause occurs at a specific embryonic stage, i.e. after formation of the germ band with cephalic lobes and telson and sequential mesoderm segmentation. As long as the eggs are incubated at 25° C, cell divisions and morphological development of the embryos cease. To examine changes in percentage of embryonic cells in the G₁, S and G₂ phases during embryogenesis, nuclear fractions were isolated from embryos, stained with propidium iodide and then subjected to flow cytometric analysis. The percentages of embryonic cells in G₁, S and G₂ were 10, 35 and 55%, respectively, at the stage of formation of cephalic lobes, whilst 98% of cells were in G₂ at diapause stage. After termination of diapause by acclimation at 5° C or by a combination of chilling and HCl, cell division resumed in the embryos. During this period, the cells rapidly entered S phase through G₁ from G₂, suggesting that their G₁ phase was short. In eggs in which diapause was averted by HCl-treatment after incubation at 25° C for 20 h after oviposition, embryonic development proceeded continuously for 9.5 days at 25° C until hatching. Along with this development, the G₁ fraction increased to levels of about 90%. These results indicate that embryonic cells are arrested in G₂ at diapause and suggest that, concomitant with further embryonic development, cell cycles become slower in proportion to an increasing length of G₁. Finally, most of the cells may be arrested in G₁, while there is only a small fraction of cells continuously cycling. Offprint requests to: T. Yaginuma     

Cold hardiness of larvae of the southwestern corn borer, Diatraea grandiosella

The cold-hardening capacity of field-collected larvae from southeast Missouri and laboratory-reared larvae of the southwestern corn borer, Diatraea grandiosella Dyar, was examined. Supercooling points of non-diapause and diapause larvae collected from maize plants grown in Missouri (36°30 N lat.) were ca.-7.0°C. The hemolymph melting points of diapause field larvae (-0.8°C) were significantly lower than those of non-diapause larvae collected in July (-0.5°C). The supercooling points of hemolymph from non-diapause and diapause field larvae ranged randomly from-10° to-18°C. Supercooling points of non-diapause laboratory larvae increased from-13° to-10°C prior to pupation, whereas those of diapause larvae increased similarly before the onset of diapause, but then decreased during diapause to ca.-17°C. No change in supercooling points or capacity to survive in the presence of ice was observed in diapause laboratory larvae acclimated at 4°C for 63 days. Laboratory and field larvae began to freeze at ca.-1.5°C in the presence of ice, but survived to several degrees below their melting points. The high supercooling points of field larvae appeared to be due to the presence of an environmental ice-nucleator. Although data for laboratory larvae indicate sufficiently low supercooling points to permit winter survival in southeastern Missouri, considerable larval mortality occurs in the field due to inoculative freezing and the presence of an ice-nucleator.     

Diapause avoidance induced by low temperature in the yellow-spotted longicorn beetle, Psacothea hilaris

The diapause-averting effect of low temperature on pre-diapause larvae was examined in the yellow-spotted longicorn beetle, Psacothea hilaris. Larvae that had been reared under diapause-inducing conditions (25 °C , L12:D12) were temporarily exposed to 10 °C for various periods, and returned to the initial condition. Diapause was not averted by chilling for 15 days irrespective of the age of the larvae at chilling. After a 30-day chilling treatment, all of the 40- and 60-day-old larvae averted diapause, while diapause was averted in only one-third of the 10- and 20-day-old larvae. None of the pre-diapause larvae chilled for 60 days entered diapause irrespective of the age at chilling. With diapause avoidance, larvae that overwintered in earlier instars can start growing in earliest spring without any arrest; this phenomenon probably subserves the synchronization of larval development in a population.     

Photoperiodic responses during larval development and diapause of two geographic ecotypes of the rice stem maggot, Chlorops oryzae

Chlorops oryzae is bivoltine in northern Japan but trivoltine in the southern part of the country. In the bivoltine strain, both the egg and larval stages were found to be sensitive to photoperiod. When the egg stage was exposed to a long-day photoperiod (16L:8D), larval development showed a short-day type response, and mature third-instar larvae entered a summer diapause under a long-day photoperiod (15L:9D). When eggs experienced short days, the first-instar larvae entered a winter diapause under short-day conditions, and the critical photoperiod in the larval stage ranged from about 14L:10D to about 12L:12D as the photoperiod experienced by the eggs increased from 12L:12D to 14L:10D. However, the development of the larvae after overwintering was not influenced by the photoperiod. In the trivoltine strain, larval development was retarded under a 14L:10D photoperiod but not under either shorter or longer photoperiods, when larvae had spent the egg stage under a 16L:8D photoperiod. The critical photoperiod of the larval stage for the induction of a winter diapause in the first instar was about 12L:12D, though it varied to some extent with the photoperiod during the egg stage. Thus, Chlorops oryzae was able to adapt itself to the local climatic conditions by the development of variable and complicated photoperiodic responses.     

Respiration of individual honeybee larvae in relation to age and ambient temperature

The CO₂ production of individual larvae of Apis mellifera carnica, which were incubated within their cells at a natural air humidity of 60–80%, was determined by an open-flow gas analyzer in relation to larval age and ambient temperature. In larvae incubated at 34 °C the amount of CO₂ produced appeared to fall only moderately from 3.89±1.57 µl mg^–1 h^–1 in 0.5-day-old larvae to 2.98±0.57 µl mg^–1 h^–1 in 3.5-day-old larvae. The decline was steeper up to an age of 5.5 days (0.95±1.15 µl mg^–1 h^–1). Our measurements show that the respiration and energy turnover of larvae younger than about 80 h is considerably lower (up to 35%) than expected from extrapolations of data determined in older larvae. The temperature dependency of CO₂ production was determined in 3.5-day-old larvae, which were incubated at temperatures varying from 18 to 38 °C in steps of 4 °C. The larvae generated 0.48±0.03 µl mg^–1 h^–1 CO₂ at 18 °C, and 3.97±0.50 µl mg^–1 h^–1 CO₂ at 38 °C. The temperature-dependent respiration rate was fitted to a logistic curve. We found that the inflection point of this curve (32.5 °C) is below the normal brood nest temperature (33–36 °C). The average Q₁₀ was 3.13, which is higher than in freshly emerged resting honeybees but similar to adult bees. This strong temperature dependency enables the bees to speed up brood development by achieving high temperatures. On the other hand, the results suggest that the strong temperature dependency forces the bees to maintain thermal homeostasis of the brood nest to avoid delayed brood development during periods of low temperature.Abbreviations m body mass - R rate of development or respiration - T_I inflexion point of a logistic (sigmoid) curve - T_L lethal temperature - T_O temperature of optimum (maximum) developmentCommunicated by G. Heldmaier     

Effects of temperature on survival, development, growth and feeding of larvae of Yellowtail clownfish Amphiprion clarkii (Pisces: Perciformes)

To understand the physiological and ecological responses of marine fishes to the change of water temperature, newly-hatched larvae of Yellowtail clownfish Amphiprion clarkii were reared in captivity at water temperatures of 23, 26 and 29 °C till they completed the metamorphosis to juvenile phase, and larval survival, development, growth and feeding were evaluated during the experimental period. The results showed that water temperature influenced the physiological performance of larvae of A. clarkii significantly. The survival and growth rates of larvae of A. clarkii increased significantly with the increase of water temperature from 23 to 29 °C (P < 0.05). Water temperature also influenced larval development of A. clarkii significantly and larvae reared at 23 °C took longer time for post-larval development and metamorphosis compared to 26 and 29 °C (P < 0.05). Total length and body weight for post-larval development and metamorphosis decreased with the increase of water temperature from 23 to 29 °C (P < 0.05). Q₁₀ in developmental rate was higher than in daily growth rate at the same rearing temperature, indicating that at water temperature had greater influence on larval development than on growth. Water temperature also influenced larval feeding of A. clarkii significantly with feed ration (FR) and feed conversion efficiency (FCE) increased with the increase of water temperature from 23 to 29 °C (P < 0.05). There was a positive correlation between FR and specific growth rate (SGR) (P < 0.05) but not between FCE and SGR (P > 0.05), indicating that FR influenced growth rate significantly in larvae of A. clarkii. This study demonstrated that the physiological responses of larvae of A. clarkii to the change of water temperature and confirmed that water temperature influenced larval survival, development, growth and feeding significantly. This study suggests that the decline of larval survival and growth rates, extension of pelagic larval duration and reduction of larval feeding at lower temperature have ecological impacts on larval dispersal and metamorphosis, juvenile settlement and population replenishment in A. clarkii in the wild.     

The effect of host age ofLymantria dispar larvae [Lep.: Lymantriidae] on the development ofGlyptapanteles liparidis [Hym.: Braconidae]

The endoparasitic development ofG. liparidis was examined in 3 different host stages of gypsy moth larvae. Hatching ofG. liparidis-larvae occurred 3 to 5 days after oviposition in hosts parasitized during their premoulting period, and after 5 to 7 days in those parasitized in the 3^rd midinstar state. The parasites generally moulted to the 2^nd larval instar between the 11^th and 13^th day in the first group, and between the 13^th and 15^th day in the latter, when they had reached a volume of 0.04–0.05 mm³. The positive correlation between host ecdysis and the ecdysis of 1^st stadium larvae to L₂ suggested that host moulting influenced the development of the parasitoid larvae. Emergence from the host larvae occurred at 20°C after 27 days on average, and coincided with the parasites moulting to the 3^rd instar. Five to 7 days after spinning their cocoons near the developmentally arrested host larva, the male, and 1 to 2 days later the female wasps eclosed. Due to the variation in the number of parasites per host, no difference was observed between the hosts parasitized at various stages; however, a tendency for later parasitized hosts to contain more parasite larvae was evident. The nutritional conditions of the moth parental generation influenced both host and parasite development. On the other hand no influence of host age was observed on emergence dates of larvae and wasps.      

Diapause induction in Ephestia cautella: An interaction between genotype and crowding

Larval diapause induction in Ephestia cautella (Walker) (Lepidoptera: Pyralidae) is a function of the interaction between genotype and larval crowding. A diapause stock in which 79% of the larvae diapaused under crowded conditions and 40% diapaused under uncrowded conditions was maintained by selection. Outbreeding of adults from this diapause stock to those from a non-diapause stock resulted in 36% diapause under crowded conditions and 6% diapause under uncrowded conditions. The rate of termination of larval diapause is inheritable and temperature dependent. These data seem to explain the seasonal trends in percentage larval diapause among E. cautella infesting citrus pulp during storage.Larval diapause was induced in groups of larvae by crowding in mass cultures and in single larvae by rearing on a small amount of fresh diet or on a larger amount of fresh diet containing residual diet from crowded cultures. The diapause-inducing effects of this residual diet could be removed by extraction with lipid solvents. Some activity was demonstrated when the extract was dried onto fresh diet.

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Resume L'induction de la diapause larvaire d'Ephestia cautella (Walker) dépend du génotype et de la densité larvaire.Des croisements à l'intérieur de la souche diapausante donnent 79% de diapause aux fortes densités larvaires et 40% aux faibles densités. Les croisements des adultes de cette souche avec ceux de la souche non-diapausante donnent 36% de diapause aux fortes densités et 6% aux faibles densités. La fréquence de fin de diapause est héréditaire et dépend de la température. Ces résultats peuvent expliquer les variations saisonnières du taux de diapause E. cautella dans la pulpe de citron stockée.La diapause larvaire a été induite chez des groupes de chenilles par surpeuplement dans des élevages standards, et chez des chenilles isolées par élevage sur une quantité limitée d'aliments frais ou sur une quantité abondante d'aliments frais contenant des résidus alimentaires provenent d'élevages surpeuplés. Les effets inducteurs de ces résidus alimentaires disparaissent après extraction avec les solvants de lipides. Une certaine action est observée par de l'extraint sec sur de l'aliment frais.

     

Termination of the facultative diapause in the codling moth, Laspeyresia pomonella (Lepidoptera, Tortricidae)

The abiotic factors regulating the termination of the facultative diapause of the mature larva of the codling moth, Laspeyresia pomonella (L.), are described. The termination of diapause under long-day conditions (LD) is influenced by: (i) the rearing temperature of the larvae during prediapause development, (ii) the duration of the preincubation period, i.e. the time for which the diapausing larvae remain under prediapause rearing conditions, (iii) the reactivating incubation, i.e. the period for which the larvae are chilled, and (iv) the complementary incubation (LD and, except in one experiment, 26°), i.e. the period after the chilling needed for the pupation of the insects. The complementary incubation is distinctly shorter if the prediapause development of the larvae takes place at rearing temperatures below 26°, i.e. 21° or 19°. The latter conditions led to 100% pupae and pupation could be accelerated by prolonging the preincubation period. In larvae reared at 26°, the prolongation of the preincubation period raised the rate of pupation to a maximum of only 56% and also caused higher mortality. On the other hand, diapause was terminated under short-day conditions if the temperature was raised to 26° after a rearing temperature of 19° and a preincubation period of 90 days.

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Beendigung der fakultativen diapause des apfelwicklers, Laspeyresia Pomonella (Lepidoptera, Tortricidae)

Zusammenfassung Es wurde untersucht, welche abiotischen Faktoren die Beendigung der fakultativen Diapause ausgewachsener Apfelwicklerlarven (Laspeyresia pomonella) steuern. Die Beendigung der Diapause unter Langtagbedingungen (LT) wird beeinflusst durch: 1. die Zuchttemperatur der Larven während der Prädiapauseentwicklung; 2. die Dauer der Präinkubation, d.h. die Zeit, während der diapausierende Larven unter Prädiapause-Zuchtbedingungen bleiben; 3. die Reaktivierungs-inkubation, d.h. eine Periode von 70 d, während der die Larven auf 4° gekühlt werden und 4. die Komplementärinkubation (LT und, ausser in einem Experiment, 26°), d.h. die nach der Kühlung bzw. dem Wechsel in der Photoperiode benötigte Zeitdauer bis zur Verpuppung. Die Komplementärinkubation ist deutlich kürzer, wenn die Prädiapauseentwicklung der Larven bei Temperaturen unter 26° stattfindet. Bei 19° verpuppten sich 100% der Larven, wobei der Zeitpunkt der Verpuppung durch die Verlängerung der Präinkubationszeit beschleunigt wurde. Bei Larven, die bei 26° gezüchtet wurden, erreichte die Verpuppungsrate lediglich 56% bei einer relativ langen Präinkubationsdauer, wobei deren Verlängerung auch die Larvenmortalität erhöhte. Nach einer Zuchttemperatur von 19° und einer Präinkubationsdauer von 90 Tagen konnte die Diapause unter Kurztagbedingungen beendet werden, wenn die Temperatur auf 26° erhöht wurde.

     

Role of photoperiod and temperature in the induction of overwintering pupal diapause in the spotted tentiform leafminer,Phyllonorycter blancardella

The role of photoperiod and temperature in the induction of overwintering diapause inPhyllonorycter blancardella (F.) (Lepidoptera: Gracillariidae) was examined in the laboratory and field using leafminers from commercial apple orchards in Ontario, Canada.P. blancardella exhibited a long-day response to photoperiod: long daylengths resulted in uninterrupted development whereas short daylengths induced diapause. The estimated critical photoperiod for diapause induction was L14.25∶D9.75. The larvae of leafminers destined to enter diapause took ca. 3× longer to complete development than the larvae of non-diapausing leafminers. The development prolonging effect of photoperiod decreased with decreasing daylength. Temperature modified the diapause inducing effect of photoperiod. At L14.25∶D9.75, diapause incidence was similar at 15 and 20°C but was lower at 25°C. Photoperiod also altered the normal relationship between development rate and temperature. At L14.25∶D9.75, the duration of larval development of diapausing leafminers was similar at 15, 20 and 25°C. Temperature alone is unlikely to have a role in the induction of diapause because leafminers exposed to natural late summer and fall temperature regimes and L16∶D8 did not enter diapause.     

Diapause termination and thermal requirements for postdiapause development inAphelinus mali at constant and fluctuating temperatures

Diapause termination under natural and simulated overwintering conditions, the effect of subzero temperature on postdiapause development and the relationship between postdiapause development rate and constant and fluctuating temperatures was studied in a Dutch population ofAphelinus mali Hald. (Hymenoptera: Aphelinidae).The rate of diapause termination was similar in larvae overwintering under natural and simulated conditions. Most larvae had terminated diapause by the last week of February. Some female larvae may have remained in diapause until the end of March. The exposure of postdiapause larvae to –10°C for two weeks did not affect their survival or postdiapause development rate.PostdiapauseA. mali larvae could complete development and the adults emerge from their mummified aphid hosts at constant temperatures from 12 to 24°C. Although some larvae completed postdiapause development at 10°C, few emerged. The theoretical threshold temperature (t_o) for postdiapause development was 9.4°C and the thermal constant (K) 136.4 degree-days. K was 121.4 and 134.8 for first and 50% emergence, respectively.The number of heat units accumulating above 9.4°C to 1st and 50% emergence was similar under constant and fluctuating temperatures.

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Fin de la diapause et exigences thermiques pour le développement après la diapause d'Aphelinus mali soumis à des températures constantes ou à des thermopériodes

Résumé L'achèvement de la diapause en conditions naturelles ou simulant l'hiver, les effets des températures inférieures à zéro sur le développement après la diapause et les relations entre la vitesse de développement après la diapause et les températures constantes ou en thermopériodes ont été examinés sur des populations néerlandaises d'A. mali (Hymenop.; Aphélinidae).Les taux d'achèvement de la diapause de larves hivernantes étaient semblables en conditions naturelles ou simulées. La plupart des larves ont terminé leur diapause la dernière semaine de février. Quelques larves femelles sont restées en diapause jusqu'à fin mars. L'exposition pendant 2 semaines des larves sorties de diapause à –10 °C ne compromet pas leur survie ou leur taux de développement après la diapause.Les larves ayant diapause peuvent terminer leur développement et les adultes émerger des pucerons momifiés aux températures constantes comprises entre 12 et 24 °C. Bien que quelques larves achèvent leur développement à 10 °C, peu émergent. La température seuil théorique de développement après la diapause (t_o) a été de 9,4 °C et la constante thermique (K), 136,5 degrés-jours. Pour la première émergence et pour 50% d'émergences, les valeurs de K étaient respectivement: 121,4 et 134,8.Le nombre d'unités thermiques pour la première émergence et pour 50% d'émergences était le même à température constante ou avec une thermopériode.

     

The response of development rate to temperature in the univoltine cranefly,Tipula subnodicornis Zetterstedt

Summary The cranefly, Tipula subnodicornis, emerges as an adult in the spring and has an annual life-cycle in the British Isles. This is maintained partly through the presence of a winter diapause but the response of development rate to temperature also acts to preserve the timing of the cycle. During development under constant temperature conditions in the laboratory the optimum temperature (taken as the temperature which promoted the most rapid development) dropped from 25°C, or above, in the egg stage to below 20°C in the late larval stages. It is suggested that at the warmer, southern limits of the geographical range rapid early development may be compensated by a retardation in late larval growth. In addition, the response of growth rate to change in temperature was small in the fourth, final, instar and resulted in low Q ₁₀ values; 2.4 between 7° and 10°C, 1.5 between 10° and 15°C and 0.9 between 15° and 20°C. As the fourth instar comprises the greater part of the growth period, this has the effect of minimising the effect of temperature differences which are the result of differences of latitude or altitude. Even at optimum temperatures the growth period was prolonged and larvae in the field do not reach maximum weight, and the photosensitive stage, until late autum when short daylength promotes diapause. Subsequent development in the spring, before pupation and during the pupal period, showed a reversion to the higher Q ₁₀ figures of the early stages in development.The development of final instar Tipula subnodicornis larvae is contrasted with that of Tipula melanoceros. Tipula melanoceros emerges as an adult in September and it is likely that it has an egg diapause. Consequently larval development is confined to a short period between April and late July and growth must be rapid during this period. Under constant temperature conditions in the laboratory the growth of final instar larvae showed a marked contrast to that of Tipula subnodicornis in that the response to temperature was large and remained positive over a wider temperature range.     

Lipid peroxidation and superoxide dismutase activity in relation to photoinhibition induced by chilling in moderate light

The effect of a chilling stress, at a moderate photon flux density for a few hours, on the peroxidation of membrane lipids and on superoxide dismutase (SOD) activity was compared in leaf slices of chilling-sensitive and chilling-insensitive plants. The aim was to determine if susceptibility to chill-temperature photoinhibition could be related to either damage to membrane lipids by superoxide and-or a decrease in activity of chloroplast SOD. Plants used were Nerium oleander L., grown at 45° C, and Cucumis sativus L., both susceptible to chill-temperature photoinhibition, and N. oleander, grown at 20° C and Spinacia oleracea L., both insensitive to chill-temperature photoinhibition. Lipid peroxidation was assessed by measuring the concentration of malondialdehyde (MDA). Leaf slices from all plants showed a basal level of MDA which decreased by about 15% when the leaf slices were chilled in the light. The level of MDA was not increased by the addition of either KHCO₃ or methyl viologen during chilling but it was increased, up to threefold, by the addition of Rose Bengal, which produces singlet oxygen. Chloroplast SOD activity was assessed in leaf extracts as the cyanide-sensitive production of H₂O₂ in a system which produced superoxide. Activity of SOD was similar in all the plants and was altered little by chilling. The results show that for the plants tested, chilling at a moderate photon flux density for 5 h does not increase the susceptibility of cell membranes to peroxidative damage nor does it decrease the activity of SOD. It was concluded that the susceptibility of chilling-sensitive plants to chill-temperature photoinhibition cannot be explained on the basis of differences in the vulnerability of membrane lipids to damage by superoxide or differences in SOD activity.Abbreviations Chl chlorophyll - MDA malondialdehyde - MV methyl viologen - O ₂ ^- superoxide - 20°-oleander Nerium oleander grown at 20° C - 45°-oleander N. oleander grown at 45° C - PFD photon flux density - SOD superoxide dismutase Deceased     

Diapause pupal color diphenism induced by temperature and humidity conditions in Byasa alcinous (Lepidoptera: Papilionidae)

We investigated whether diapause pupae of Byasa alcinous exhibit pupal color diphenism (or polyphenism) similar to the diapause pupal color polyphenism shown by Papilio xuthus. All diapause pupae of B. alcinous observed in the field during winter showed pupal coloration of a dark-brown type. When larvae were reared and allowed to reach pupation under short-day conditions at 18 °C under a 60 ± 5% relative humidity, diapause pupae exhibited pupal color types of brown (33%), light-brown (25%), yellowish-brown (21%), diapause light-yellow (14%) and diapause yellow (7%). When mature larvae reared at 18 °C were transferred and allowed to reach pupation at 10 °C and 25 °C under a 60 ± 5% relative humidity after a gut purge, the developmental ratio of brown and light-brown, yellowish-brown, and diapause light-yellow and diapause yellow types was 91.2, 8.8 and 0.0% at 10 °C, and 12.2, 48.8 and 39.0% at 25 °C, respectively. On the other hand, when mature larvae reared at 18 °C were transferred and allowed to reach pupation at 10 °C, 18 °C and 25 °C under an over 90% relative humidity after a gut purge, the developmental ratio of brown and light-brown, yellowish-brown, and diapause light-yellow and diapause yellow types was 79.8, 16.9 and 3.3% at 10 °C, 14.5, 26.9 and 58.6% at 18 °C, and 8.3, 21.2 and 70.5% at 25 °C, respectively. These results indicate that diapause pupae of brown types are induced by lower temperature and humidity conditions, whereas yellow types are induced by higher temperature and humidity conditions. The findings of this study show that diapause pupae of B. alcinous exhibit pupal color diphenism comprising brown and diapause yellow types, and suggest that temperature and humidity experienced after a gut purge are the main factors that affect the diapause pupal coloration of B. alcinous as environmental cues.     

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

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

Effects of temperature on larval eclosion of the winter moth, Operophtera brumata

The developmental threshold and heat-unit requirements for larval eclosion of the winter moth, Operophtera brumata (L.) [Lepidoptera: Geometridae], were determined from experiments involving eggs exposed to various chilling and warming treatments. The developmental threshold was determined to be 4 °C. Eggs which were chilled below the developmental threshold required fewer heat-units for larval eclosion than did eggs not given a chill treatment. Eggs cultured at 14 °C required 470 (±47) degree-days for 50% larval eclosion. However, eggs chilled for 2 weeks at 1 °C and subsequently placed at 14 °C required 382 (±33) degree-days while eggs chilled for 12 weeks at 1 °C and subsequently placed at 14 °C required 156 (±12) degree-days. The results are discussed in relation to chilling intensity, diapause, and physiological-time models.

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Résumé Le seuil de développement et la quantité de chaleur nécessaires pour l'éclosion larvaire chez Operophtera brumata (L.) (Lepidoptera: Geometridae) ont été déterminés en exposant les oeufs à différents régimes de refroidissement suivis de périodes de réchauffement. Le seuil de développement déterminé est de 4 °C. Les oeufs placés d'abord à des températures inférieures à celle du seuil de développement demandent moins de quantité de chaleur pour l'éclosion larvaire, que les oeufs qui n'ont pas été exposés à de basses températures. Les oeufs maintenus à 14 °C demandent 470 (±47) degrés-jours pour permettre 50% d'éclosion. Cependant, des oeufs maintenus pendant 2 semaines à 1 °C, puis à 14 °C, exigent 382 (±33) degrés-jours; ceux placés pendant 12 semaines à 1 °C, puis à 14 °C, en exigent 156 (±12). Les résultats sont discutés en relation avec l'intensité du refroidissement, la diapause et les modèles physiologico-temporels.