Effect of photoperiod on growth and gonadal development of juvenile Topmouth Gudgeon Pseudorasbora parva

In this study, we investigated the effect of photoperiod on growth and gonadal development of juvenile topmouth gudgeon, Pseudorasbora parva. Seven different photoperiods (light:dark, L:D) were tested: 0L:24D, 4L:20D, 8L:16D, 12L:12D, 16L:8D, 20L:4D and 24L:0D, from November to January. Throughout the study, light intensity was kept at 400 lx on the water surface, and the fish were fed a commercial diet twice a day. Weight gain, feed intake, specific growth rate, and feed conversion efficiency were significantly higher under the constant–long day photoperiods (24L:0D, 20L:4D and 16L:8D) than those under the short–no day photoperiods (12L:12D, 8L:16D, 4L:20D, 0L:24D) (p?<?0.05). The highest mean female gonadosomatic index (GSI) and mean oocyte size were observed under 24L:0D followed by 20L:4D and 16L:8D photoperiods, and these parameters were significantly higher than those of fish cultured under the short–no day photoperiod regimes (p?<?0.05). Proportion of females with GSI?>?9 % were 50.00 %, 46.67 %, 48.48 %, in 24L:0D, 20L:4D and 16L:8D, respectively. These results demonstrate that the growth of juvenile P. parva can be stimulated significantly by constant–long photoperiods and that these photoperiods can advance sexual maturity of females by approximately 2 months.     

Photoperiod and temperature studies to determine whether diapause is found in successive generations of the African phytoseiid,Euseius fustis (Pritchard and Baker) (Acari: Phytoseiidae)

Photoperiod and temperature conditions known to induce diapause in tropical arthropods were tested on two generations (G₀ and G₁) of the phytoseiid, Euseius fustis. Failure to lay eggs or a pre-oviposition period longer than 15 days were the criteria used to determine whether females were in diapause. Females reared from egg to adult and held throughout adult life under cyclic temperatures of 29/20°C in combination with long photophases of 16L: 8D and 14L: 10D showed no indication of aestival diapause. Similarly, hibernal diapause was not induced in females reared under a constant temperature of 18°C and a photophase of 8L: 16D. Under the various test conditions, females initiated oviposition within an average of 4 days. Overall, pre-oviposition patterns for G₀ and G₁ females were similar under the same test conditions. Reproductive patterns based on the mean number of eggs per female per day varied only slightly between generations for the same treatments. No behavioural or morphological attributes associated with diapause were observed.     

Photoperiod- and temperature-dependent induction of larval diapause in a multivoltine bruchid, Bruchidius dorsalis

The wild bruchid beetle, Bruchidius dorsalis Fahraeus (Coleoptera: Bruchidae), has a multivoltine life cycle and overwinters in several developmental stages in the middle part of Japan. We investigated the incidence of diapause under different conditions of photoperiod (from L8:D16 to L16:D8) and temperature (at 20 °C and 24 °C). Our experiments revealed the following results: (1) B. dorsalis entered diapause at the final (late fourth) instar larva under short photoperiods, (2) the larval diapause incidence was dependent on temperature (critical photoperiods were 12.5 h at 20 °C and 12 h at 24 °C), (3) some individuals did not enter diapause under short-photoperiod conditions at 24 °C, and (4) the sensitive stages to the photoperiod were from the late egg stage to the early first instar larva. Based on these results, we discuss not only the evolution of a complex overwintering strategy inB. dorsalis but also the domestication process of stored-bean pests.     

Exposure of <Emphasis Type="Italic">Eocanthecona furcellata</Emphasis> (Hemiptera: Pentatomidae) nymphs and adults to high temperatures induces an aestivo-hibernal egg diapause: a strategy for surviving hot summers

The predatory stink bug Eocanthecona furcellata (Wolff) (Hemiptera: Pentatomidae), which is widely distributed in the subtropics and tropics, is a potential biological control agent for lepidopteran, coleopteran, and hemipteran pests. We investigated development of eggs oviposited by E. furcellata females subjected to various photoperiods (i.e., 8L:16D, 12L:12D, and 16L:8D) and temperature regimes (i.e., 26:26, 30:28, 32:30, 34:32, 35:33, and 36:34 °C; temperatures at light and dark phases, respectively) at different developmental stages. Exposure of E. furcellata nymphs and adults to different photoperiods and temperature regimes affected the development of eggs they subsequently oviposited. Temperatures greater than 30 °C induced diapause in a portion of the eggs, as evidenced by the delayed appearance of the black circle and the red spot, characteristics of early and late stage of embryonic development, respectively. The proportion of diapausing eggs generally increased with increasing temperature exposure, whereas it decreased with advancing developmental stage/age at which E. furcellata was subjected to the temperature regime. Although viability of diapausing eggs was lower than that of non-diapausing eggs, diapausing eggs survived longer under high temperatures. The trade-off between survivability and viability may be a strategy to survive unpredictability of summer temperatures.     

Induction and termination of diapause inOrius predatory bugs

Photoperiodic induction of reproductive diapause at 18°C was investigated in fourOrius [Heteroptera: Anthocoridae] species.Orius insidiosus (Say) displayed a long-day response with a critical photoperiod between L11:D13 and L12:D12. Diapause in this species was terminated rapidly when the temperature and/or the daylength were increased.Orius majusculus (Reuter) also displayed a long-day response. The critical photoperiod fell between L14:D10 and L16:D8. Diapause in this species was not terminated within 14 days when both temperature and daylength were increased. InOrius albidipennis (Reuter) no diapause could be induced at photoperiods varying from L8:D16 to L16:D8. InOrius tristicolor (White) a high proportion of diapause was found at all photoperiods tested. The effect of temperature on photoperiodic induction of diapause was studied inO. insidiosus at L10:D14. Diapause occurred at 18°C, 21°C and 25°C, but not at 30°C. Again, diapause was terminated rapidly after transfer to 25°C/L16:D8. Exposing only the nymphal instars 1–5 to short daylength was not enough to induce diapause in the whole population ofO. majusculus. Orius predatory bugs are used as biocontrol agents against western flower thrips,Frankliniella occidentalis (Pergande) [Thysanoptera: Thripidael, in greenhouses. The consequences of photoperiodic induction of diapause for the success of early season releases ofOrius are discussed.     

PHOTOPERIODIC CONTROL OF ADULT DIAPAUSE IN CHRYSOPERLA SINICA (TJEDER) (NEUROPTERA: CHRYSOPIDAE— I. CRITICAL PHOTOPERIOD AND SENSITIVE STAGES OF ADULT DIAPAUSE INDUCTION

Abstract  Photoperiodic sensitivity for diapause induction of the green lacewing, Chrysoperla sinica (Tjeder) was examined at 22°C. The adult diapause of C. sinica was induced by short-day photoperiods, and the critical photoperiod for its induction was between 12.5L-11.5D and 13L-11D.
Adults developed without diapause under long-day conditions, and entered diapause under short-day conditions. The adult stage was the uppermost sensitive stage for adult diapause induction, adults could go into diapause only when the emerging adults were under diapause-inducing short-day photoperiods. The short-day photoperiodic experience by transferring between 15L: 9D and 9L: 15D at preimaginal stages did not result in adult diapause under 15L: 9D photo regime, although some treatments extended the pre-oviposition period in adult stage. The results showed that the 3rd instar larvae and pre-pupae were more sensitive to the photoperiodic change from 15L: 9D to 9L: 15D photo regime than the other preimaginal stages.     

Control mechanisms of larval diapause and quiescence in Trogus mactator, an endoparasite of papilionids

First instar larvae of Trogus mactator Tosquinet (Hymenoptera: Ichneumonidae) developed to second instar immediately after pupation of the host, Papilio xuthus L., under a short photoperiod (L10: D14) as well as under a long photoperiod (L15: D9). The host went into diapause as pupa under short photoperiods, and the parasite at the third instar in the host pupa. They responded independently to short photoperiods. The critical photoperiods for diapause induction both in the parasite and the host were between 12 and 13 h, but there was a slight disparity between them. Diapause of neither the parasite nor the parasitized host was terminated under short photoperiods. Diapause of the parasite in a diapausing host pupa was completed when it was transferred to an incubator having a long photoperiod. This suggests that the exposure to a long photoperiod breaks diapause in the parasite irrespective of physiological conditions of the host. The period from the change in photoperiod to time of parasite emergence shortened as the change was made later. The parasite remaining in a host pupa for 60 days completed diapause even under a short photoperiod when it was implanted into a nondiapausing host pupa, or when 20-hydroxyecdysone was injected into the parasitized and diapausing host pupa. These results indicate that the physiological development for diapause termination in T. mactator seems to complete 60 days after host pupation and endogenous ecdysteroids are somehow involved in termination of diapause in the parasite.

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Résumé Que la photopériode soit brève (L10/D14) ou longue (L15/D9), le premier stade de T. mactator Tosquinet (Hym. Ichneumonidae) donne un second stade immédiatement après la nymphose de son hôte, Papilio xuthus L.,. L'hôte entre en diapause au stade nymphal sous photopériodes courtest et le parasite au troisiéme stade larvaire dans la nymphe de l'hôte. Ils réagissent indépendamment aux courtes photopériodes. Les photopériodes critiques pour l'induction de la diapause, tant de l'hôte que du parasite, se situent entre 12 et 13 h, mais il y a une légère différence. Ni la diapause de l'hôte, ni celle du parasite ne se terminent sous de courtes photopériodes. Le transfert dans une étuve ayant une longue photopériode rompt la diapause du parasite dans un hôte diapausant. Ceci laisse supposer que l'exposition à une longue photopériode rompt la diapause du parasite quel que soit l'état physiologique de l'hôte. Le délai entre le moment du changement de la photopériode et la date d'émergence du parasite, est d'autant plus raccourci que le changement est tardif. Les parasites restés dans une nymphe hôte pendant 60 jours, achèvent leur diapause même avec une photopériode courte quand ils sont transplantés dans un hôte sans diapause, ou quand de la 20-hydroxyecdysone a été injectée dans la nymphe hôte parasitée et diapausante. Ces résultats montrent que le développement physiologique nécessaire à la fin de la diapause semble achevé 60 jours après la nymphose de l'hôte et que les ecdystéroïdes endogènes semblent impliqués d'une manière ou d'une autre dans la fin de la diapause du parasite.

     

Cloning and expression patterns of the brine shrimp (Artemia sinica) glycogen phosphorylase (GPase) gene during development and in response to temperature stress

Glycogen serves as a metabolic reserve and is involved in macromolecular synthesis. Glycogen phosphorylase (GPase) is a key enzyme involved in intracellular glycogen catabolism, catalyzing the first step in glycogen degradation. In the diapause, GPase catalyzes glycogen into the closely related molecule, sorbitol. In this study, the full-length cDNA of the GPase gene (2,790 bp) was isolated from Artemia sinica for the first time by rapid amplification of cDNA ends technology. The GPase gene encoded a protein of 853 amino acids belonging to the Glycosyltransferase GTB type superfamily. The expression pattern and location of GPase were investigated at various stages during the embryonic development of A. sinica using real-time PCR and in situ hybridization. High GPase expression was detected at the 0 and 5 h stages. Subsequently, expression declined and was maintained at a low level during the stages from 10 to 40 h following by a small increase at day 3. Expression was downregulated at temperatures ranging from 25 to 20 °C and was subsequently upregulated in the range 15–5 °C. In situ hybridization assays showed wide distribution of the GPase gene during different developmental stages. From the results of this study, we conclude that the GPase gene expression is stress-related and might play an important role in Artemia development and metabolism.     

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.     

Effect of photoperiod on the induction and maintenance of diapause and on development of the predatory bugPodisus maculiventris (Hem.: Pentatomidae)

The predatory bugPodisus maculiventris Say displays a reproductive diapause. In the laboratory, at 23±1°C, when the pre-imaginal stages were reared on larvae ofGalleria melonella L. (Lepidoptera: Pyralidae) under a short photoperiod of 8L: 16D, diapause was induced in all adult females, whereas under a long photoperiod of 16L: 8D none entered diapause. Nymphal development was faster under the diapause-inducing short photoperiod than under the diapause-averting long photo-period. By contrast, embryonic development was faster under the long than under the short photoperiod. Diapause maintenance was also under the control of photoperiod. After a 10-d-period of chilling at 4°C, diapause was quickly terminated in adults kept under long photoperiod whereas it was maintained in most of those kept under short photoperiod. The predation rate of non-diapause or post-diapause adult females was much higher than that of diapause females.     

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.     

Photoperiodic responses of pupa and adult determining maternal influence on progeny diapause in two species of the genus Trichogramma (Hymenoptera, Trichogrammatidae)

The adult photoperiodic responses of two Trichogramma species were investigated by exposing adults during 3 days to one of the seven light: dark regimes: L: D = 4: 20, 8: 16, 12: 12, 14: 10, 16: 8, 18: 6, and 20: 4. The preimaginal stages of these individuals developed under short (L: D = 12: 12), long (L: D = 20: 4) or intermediate photoperiods (L: D = 14: 10 and L: D = 16: 8 for Trichogramma principium and T. embryophagum, respectively). The progeny of these females developed under short day L: D = 12: 12 and at two moderately diapause-inducing temperatures (13 and 14°C for T. principium, 14 and 15°C for T. embryophagum). In both Trichogramma species which developed at both temperatures, the percentage of diapausing prepupae was significantly dependent both on the photoperiodic conditions of the preimaginal development of the maternal generation and on the photoperiod which influenced the adult females. The adults showed a typical long-day photoperiodic response with a threshold day length of ca 13 h in T. principium and ca 16 h in T. embryophagum, which practically coincided with the thresholds of the pupal photoperiodic responses of these species revealed in our previous studies. However, the ultra-short photoperiods (L: D = 4: 20 and 8:16) caused a relatively stronger diapause-inducing effect on the progeny when applied to the adult females than when it was applied to the pupae. Thus, in both the Trichogramma species studied, the patterns of photoperiodic responses of pupa and adult were somewhat different although they almost coincided in the “ecologically significant” part of the photoperiodic scale.     

Crowding inhibits pupation inTribolium freemani: contact chemical and mechanical stimuli are involved

The relationship between the development ofCoccinella septempunctata brucki Mulsant (Coleoptera: Coccinellidae) and its parasitoid,Perilitus coccinellae (Schrank) (Hymenoptera: Braconidae) was studied at two photoperiods (L 16:D8 and L 12: D12) at 26°C. The development ofP. coccinellae is well synchronized with the physiological state of the host,C. septempunctata, which can be parasitized not only as adult but also as larva or pupa. The parasitoid larva completed larval development within 19 days in a non-diapausing host, while in diapausing adults as well as in pupae held at diapause-averting conditions, the parasitoid larva ceased growth at the first instar. Growth was resumed when diapause of the host terminated or by the emergence of the adult host from the pupa. About 550 spheric cells, teratocytes, were liberated into the host hemocoel when the parasitoid egg hatched. The teratocytes increased in size in the active host, while their development was arrested in the diapausing host. Application of methoprene caused diapause termination of both host and parasitoid larva. The results indicate that the development of the larva ofP. coccinellae depends on the physiological conditions of the host,C. septempunctata brucki. The host-parasite relation thus represents an ‘endogenous synchronization’ in the sense of Schoonhoven's definition.     

Photoperiodic induction of the pupal diapause in the tobacco hornworm, Manduca sexta

A study was made of photoperiodic induction of the facultative pupal diapause in the tobacco hornworm, Manduca sexta, reared on artificial diet in the laboratory. The species entered a prolonged diapause when the egg and larval feeding stages were reared in daily photoperiods of 13·5 hr or less. Diapause was induced in all insects at photoperiods ranging from 1 to 13 hr, and part of the population entered diapause at only 15 to 30 min of light per day. Photoperiods of 14 hr or more and continous darkness prevented diapause. Duration of diapause varied with the inductive photoperiod in which the hornworms were reared during the sensitive period. Insects reared in longer diapause-inducing photoperiods within a range of 12 to 13·25 hr remained in diapause longer than those reared in shorter photoperiods. There was no difference in the rate of larval development of hornworms reared in diapause-inducing vs diapause-preventing photoperiods. Temperatures of 26 to 30°C were most favourable for the photoperiodic induction of diapause; at 21°C, the critical photoperiod and incidence of diapause were decreased. Diapause induction was suppressed by low (18°C) and higher (33°C) temperatures. The number of inductive 12L:12D (light = 12 hr; dark = 12 hr) cycles required to induce diapause ranged from as few as 5 for some insects to as many as 12 for others when the post-inductive régimen was continuous light, but with insects previously held in continuous dark, as few as 2 12L:12D cycles during the last 2 days of larval feeding induced diapause in 38 per cent of the population. Only 3 to 4 cycles of 15L:9D during the final larval instar reversed inductive effects of 14 to 15 12L:12D cycles. Photoperiodic sensitivity extended from the late embryo to the end of larval feeding but showed considerable fluctuation during development with maximum sensitivity occurring just before egg hatch and during larval growth.Light breaks applied at different times during the dark period of 12L:12D cycles generated different response curves, depending on the number of cycles in which light breaks were repeated. When repeated for 6 cycles, a unimodal response curve was obtained; 10 cycles produced a bimodal curve and light breaks given for 18 cycles throughout the sensitive period averted diapause regardless of time of night applied. It is suggested that diapause is regulated by a photo- and thermolabile substance that accumulates during long nights (11 hr or more) and acts during the early pupal stage to inhibit the translocation and release of development-promoting neurosecretion from the brain.     

Environmental control of the seasonal life cycle of a zoophytophagous mirid, <Emphasis Type="Italic">Adelphocoris triannulatus</Emphasis> (Hemiptera: Miridae)

Adelphocoris triannulatus (Stål) (Heteroptera: Miridae) is a zoophytophagous bug that inhabits grasslands and crop fields, and is widely distributed in eastern Asia. In this study, the seasonal occurrence and environmental factors controlling the seasonal life cycle of this bug were investigated in a local population in southern Kyushu, Japan. When insects were reared under different photoperiods at 20 and 25 °C, they produced diapause or nondiapause eggs depending on the photoperiod: when reared under short-day photoperiods (≤?13 h/day), females laid diapause eggs, whereas they laid nondiapause eggs when reared under long-day photoperiods (≥?14 h/day). Because the developmental duration of the pre-adult stages and maturation (minimum duration of one generation) is estimated to be approximately 30 days at summer temperatures, these results suggest that this bug has a multivoltine seasonal life cycle. In agreement with the laboratory results, only nondiapause eggs were produced until mid-August in the field, and an increasing proportion of diapause eggs were produced from early September. In addition to these findings, field sampling from spring to autumn in 2009 indicated that three or four generations were produced in one year. These results suggest that egg diapause plays a role in preventing nymphs of A. triannulatus from developing in winter, and thereby ensures the survival of the population in adverse seasons.     

Inheritance of the photoperiodic response controlling imaginal summer diapause in the cabbage beetle, Colaphellus bowringi

Adults of the cabbage beetle Colaphellus bowringi display a summer diapause in response to the exposure of their larvae to long photoperiods. In the present study, the inheritance of the photoperiodic response controlling summer diapause in C. bowringi by crossing a high diapause strain (D strain) with a laboratory selected nondiapause strain (N strain) was investigated under different photoperiods at 22, 25 and 28 °C. The beetles in both reciprocal crosses and backcrosses showed a clear short-day response for the induction of diapause at all temperatures, similar to that of the D strain, suggesting that photoperiodic response of this beetle is heritable. The diapause incidences in the progeny from all the crosses under LD 15:9 or LD 12:12 at 25 °C suggest that genetic and genetic-environmental interactions are involved in diapause induction. The incidence of diapauses in F₁ progeny was significantly lower than that in the D × D strain but significantly higher than that in the N × N strain, indicating that the diapause capability is inherited in an incomplete dominant manner. The incidence of diapause was greater among the offspring of hybrid females when those females had a D strain mother or grandmother rather than a N strain mother or grandmother, indicating that maternal effects on diapause induction are stronger than paternal effects. The laboratory selected nondiapause strain also showed a short-day photoperiodic response at a low temperature of 22 °C, indicating that the photoperiodic photoreceptor and photoperiodic clock still function in the nondiapause strain.     

Geographical variation in photoperiodic induction of larval diapause in the bruchid beetle, Bruchidius dorsalis: polymorphism in overwintering stages

The bruchid beetle, Bruchidius dorsalis Fahraeus (Coleoptera: Bruchidae), has a multivoltine life cycle and shows geographical variation of overwintering stages in Japan. Our previous study found that B. dorsalis enters larval diapause in the final instar under short photoperiods. In cooler areas, we observed that most individuals overwinter in the final larval stage in diapause, whereas beetles at different developmental stages (non‐diapausing young instars, diapausing instars, and adults) were overwintering in warmer areas. In this study, we investigated geographical variation in the photoperiodic response for induction of larval diapause at 20 °C (three populations) and 24 °C (two populations) to clarify the overwintering strategy of B. dorsalis. We observed that (1) diapause incidence at 20 °C changed sharply from ca. 100% to 0% with a change in photoperiod in all the populations, (2) critical photoperiod was longer at 20 °C in populations from cooler areas, and (3) critical photoperiod at 24 °C was shorter than at 20 °C and a fraction of the larvae did not enter diapause, even under short photoperiods. Overwintering stages estimated from these results were consistent with those actually observed in the field. This study indicates that the geographical variation of overwintering stages is likely to reflect adaptive diapause induction in each local environment.     

Host plant mediation of diapause induction in the cabbage beetle, Colaphellus bowringi Baly （Coleoptera： Chrysomelidae）

The cabbage beetle, Colaphellus bowringiBaly （Coleoptera： Chrysomelidae）, is a serious pest of crucifers in China, undergoing an imaginal summer and winter diapause in the soil. The effects of host plants on diapause incidence were tested in the beetle. The ratio of adults entering diapause was significantly low when they fed on the mature leaves of Chinese cabbage Shanghaiqin （Brassica chinesis var communis） than those feeding on Chinese cabbage Suzhouqin （Brassica chinesis var communis）, radish （Raphanus sativus var longipinnatus） and stem mustard （Brassicajuncea vat tumida） at 25℃ combined with 13：11 （L： D） h. Fewer adults entered diapause on young leaves compared to physiologically aged and mature radish leaves at 25℃ combined with 13：11 （L： D） h. The effect of host plant species on diapause induction was also evident under continuously dark rearing conditions or at different photoperiods. These experimental results demonstrate that host plant mediation of diapause induction exists in the cabbage beetle. However, at temperatures ≤20℃ or photoperiods of 16：8 （L： D） h combined with 25℃, all individuals entered diapause regardless of the host plants, indicating that the effects of host plants on diapause induction could be expressed only within a limited range of temperatures and photoperiods.     

Temperature-dependent development, diapause and cold tolerance of Gratiana graminea, a potential biological control agent of Solanum viarum in Florida, USA

The objectives of this study were to examine temperature-dependent development, diapause and cold tolerance of Gratiana graminea Klug (Chrysomelidae), a candidate biological control agent of tropical soda apple, Solanum viarum Dunal (Solanaceae). Immature development was examined at six constant temperatures ranging from 15°C to 30°C. Diapause induction was determined by exposing adults to either long or short photoperiods at 20°C and cold tolerance was assessed by exposing adults to 0°C. G. graminea completed development at temperatures ranging from 20°C to 30°C. Linear regression estimated a lower temperature threshold of 11.7°C and 312 degree-days were required to complete development. Diapause was induced when adults were exposed to short photoperiods (10:14 L:D h) at 20°C. The lethal times for diapausing adults of G. graminea at 0°C (LT₅₀?=?19?days, LT₉₀?=?41?days) were two times higher compared to Gratiana boliviana Spaeth, a biological control agent already established in south and central Florida, USA. The presence of diapause and the greater cold tolerance suggest that G. graminea may establish and perform better than G. boliviana in northern Florida.     

Environmental induction of larval diapause and life-history consequences of post-diapause development in the Large Copper butterfly,Lycaena dispar (Lepidoptera: Lycaenidae)

Many insects in temperate zones withstand the adverse conditions of winter through entering diapause and the two most important environmental stimuli that induce diapause are photoperiod and ambient temperature. The Large Copper butterfly, Lycaena dispar Haworth (Lepidoptera: Lycaenidae), is a Palearctic butterfly that hibernates as larvae. Since this butterfly is a near threatened species in some regions, there has been a growing need for a standardized protocol for mass rearing of this butterfly based on the adequate knowledge of its ecology. In the present study, we first identified that L. dispar larvae were sensitive to the photoperiodic induction of diapause during their first larval instar. We then investigated to what extent the diapause-inducing effects of photoperiod could be modified by ambient temperatures in L. dispar larvae by exposing them to the range of day-lengths (L:D 14:10, 12:12, 10:14 and 8:16) at three different temperatures (15, 20 and 25 °C). All larvae were induced to enter diapause at low ambient temperature (15 °C) regardless of photoperiod, whereas most of them (86 %) exhibited direct development when temperature was high (25 °C). The photoperiodic induction of diapause was evident when day-length was shorter than 14 h at intermediate temperature (20 °C). Pre-diapause development was prolonged at low temperatures. Finally, we found that post-diapause development of L. dispar larvae was determined by both the chilling temperature experienced by diapausing larvae and the duration of larval diapause. Adult emergence was enhanced when larvae were chilled at 8 °C and when they had been under the state of diapause for 20 days before they were treated to terminate diapause.