Effect of different artificial diets on diapause induction under controlled temperature and photoperiod in the silkworm, Bombyx mori L.

ABSTRACT. Embryonic diapause of the silkworm, Bombyx mori , is generally induced by temperature and photoperiod during the egg stage of the previous generation and not in the larval stage. However, when silkworm larvae are reared on an artificial diet instead of mulberry leaves, their diapause is strongly affected by temperature and photoperiod experienced in the larval stage, with a distinct long-day response for diapause induction. Moreover when larvae which have been reared on artificial diet under long-day condition are fed mulberry leaves even for a short period of time, most of the resultant female adults lay diapause eggs. These results suggest that the photoperiodic response of larvae for diapause induction may be strongly suppressed by some components in mulberry leaves.     

Induction of adult diapause and of low and high reproductive states in a parasitoid wasp,Ooencyrtus nezarae, by photoperiod and temperature

Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) was reared on eggs ofRiptortus clavatus (Thunberg) (Heteroptera: Alydidae) at various temperatures under long-day (L16:D8) or short-day (L10:D14) conditions. There was no diapause during egg, larval or pupal stages under any set of conditions examined. However, at 15°C under short-day conditions, vitellogenesis was arrested in all adult females and they entered diapause. At 15°C under long-day conditions, or at 20°C under short-day conditions, some adult females entered diapause. Under the latter set of conditions, the adult females laid eggs but they laid fewer eggs than under long-day conditions, Even at 25°C, under short-day conditions, adult females laid fewer eggs than under long-day conditions, and this low rate of oviposition was attributed to the retarded development of ovaries. Diapause adults reared at 15°C were more resistant to low temperature than nondiapause adults reared at 25°C.     

Induction of diapause in the pupa of Phalaenoides glycinae by a hormone from the suboesophageal ganglion

The pupal diapause in Phalaenoides glycinae is facultative, and diapause is induced when larvae are reared in short days at low temperatures. By varying the stimulus and especially by changing larvae from one régime to the opposite during their development it is possible to induce diapause covering a wide and continuous spectrum of intensity. Wounding tends to intensify the diapause if it already exists no matter how weakly, but superficial wounding will not put a strongly non-diapausing individual into diapause. In P. glycinae diapause is caused by a hormone secreted by the suboesophageal ganglion. It is suggested that the intensity of the diapause depends on the concentration of the hormone and the duration of the period that the target (the neurosecretory cells of the brain) remains vulnerable. It is suggested that the suboesophageal ganglion may have the function of secreting a diapause-inducing hormone quite generally in short-day and long-day insects with facultative diapause and in species with obligate diapause as well.     

Quantitative short-day photoperiodic response in larval development and its adaptive significance in an adult-overwintering cerambycid beetle, Phytoecia rufiventris

The chrysanthemum longicorn beetle, Phytoecia rufiventris, overwinters in the adult stage and reproduces in spring. Larvae of this beetle develop during summer inside a host stem or root. In the present study, photoperiodic control of larval development and its adaptive significance were examined in this beetle using an artificial diet. Larvae showed a short-day photoperiodic response at 25 °C with a critical day length of around 14 h; larvae reared under short-day conditions pupated, whereas those reared under long-day conditions entered summer diapause with some supernumerary molts and did not pupate. A similar response was found at 30 °C, but with a shorter critical day length. Below the critical day length, a shorter day length corresponded to a shorter larval period. Larvae transferred from long-day conditions to various photoperiods showed a similar quantitative response. Field rearing of larvae starting at various times of year showed that pupation occurs within a relatively short period in early autumn. Field rearing of pupae and adults at various times indicated that only pupation in early autumn results in a high survival rate until winter. Earlier or later pupation led to a low survival rate due to death before overwintering in the adult and pupal stages, respectively. Thus, in P. rufiventris, timing of pupation regulated by the quantitative short-day photoperiodic response is vital for survival. Relatively lower developmental threshold in the pupal stage supports this hypothesis.     

Artificial selection for responsiveness to photoperiodic change alters the response to stationary photoperiods in maternal induction of egg diapause in the rice leaf bug Trigonotylus caelestialium

Female adults of the rice leaf bug Trigonotylus caelestialium (Kirkaldy) (Heteroptera: Miridae) produce non-diapause eggs under long-day conditions, whereas they produce diapause eggs under short-day conditions. These egg-production modes change following a photoperiodic change from long-day to short-day conditions or vice versa, with individual variations in responsiveness shown in the time from the photoperiodic change to the mode change. Strains of this insect with higher or lower responsiveness to photoperiodic change were established after several generations of selection, indicating that the individual variation has a genetic basis. The selected strains that were more responsive and less responsive to one photoperiodic change were found to be less responsive and more responsive to the opposite photoperiodic change, respectively, indicating a significant negative correlation between responsiveness to reciprocal photoperiodic changes. The selected strains also had a significantly different incidence of diapause-egg producers in stationary photoperiods compared to a non-selected strain, showing that selections for responsiveness to photoperiodic change were essentially the same as selections for a higher or lower incidence of diapause-egg producers. These results indicate that responsiveness to photoperiodic change is one aspect of the tendency to produce diapause or non-diapause eggs.     

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.     

Summer dormancy ensures univoltinism in the predatory bug Picromerus bidens

The seasonal cycle of Picromerus bidens L. (Heteroptera: Pentatomidae) is usually considered to be univoltine with an obligatory winter egg diapause. Seasonal adaptations of the species were studied in the laboratory and in field experiments. When reared under short-day photoperiodic conditions (L12:D12 and L14:D10), all females began to lay eggs synchronously soon after their emergence. However, in the females reared under long-day conditions (L18:D6 and L20:D4) and outdoors in June–July, oviposition was significantly delayed. This delay in reproduction induced by photoperiodic conditions and then spontaneously terminated was considered to be aestivation. Egg batches laid by females in the laboratory and in the field were kept at 25 °C for two months. From 30.8 to 93.8% of batches contained eggs which hatched without cold treatment between day 14 and 60 after oviposition. The proportion of eggs hatched was 17.7 to 20.9% in the short-day regimes, while it was significantly less (5.7 to 6.0%) under long-day conditions. It is concluded that in some eggs diapause is of low intensity and that if under natural conditions the first batches had been laid at the end of June, nymphs would have hatched at least from some eggs during the same season even without cold treatment. Such untimely hatching would have resulted in the death of nymphs and adults unprepared for overwintering. A photoperiodic response which induces aestivation in the early emerging adults in June–August may prevent early oviposition and occurrence of a second generation and thus maintains univoltinism in P. bidens.     

In vivo and in vitro photoperiodic induction of diapause using isolated brain-suboesophageal ganglion complexes of the silkworm,Bombyx mori

By use of a bivoltine silkworm race which shows a long-day photoperiodic response after induction during the last (5th) instar, we tried to programme photoperiodic induction in the isolated brain-suboesophageal ganglion complex in vivo and in vitro. A pair of the complexes from a newly ecdysed 5th-instar female was transplanted into the abdomen of a late 5th-instar larva and exposed to long-day (20 h light: 4 h dark) or short-day (8 h light: 16 h dark) conditions for 3 cycles. The short-day-exposed complexes elicited the production of diapause eggs in the recipient silkworms destined to become non-diapause egg producers, whereas the long-day-exposed brain complexes produced non-diapause eggs. Transplant experiments of the brain-suboesophageal ganglion complex using isolated abdomens showed a similar result. The brain complexes from newly ecdysed females of the 5th-instar were cultured in Grace's insect medium under 20 h light: 4 h dark or 8 h light: 16 h dark for 4 cycles, respectively. After in vitro culture, a pair of complexes was implanted into the abdomen of a late 5th-instar larva destined to become a non-diapause egg producer, and the diapause incidence in the resultant moths was examined. The brain complexes which received the short-day cycles induced a large portion of diapause eggs, whereas those which received the long-day conditions induced non-diapause eggs. The connection of corpora cardiaca and corpora allata with the brain complex had no influence on the result. Suboesophageal ganglia which had been cultured in vitro and implanted elicited a remarkable production of diapause eggs, but cultured brains were ineffective in producing diapause eggs, regardless of the photoperiod experienced. These results demonstrate that photoperiodic induction of the silkworm can be programmed in in vivo and in vitro culture systems, and that components of the photoperiodic clock (photoreceptor, clock, and counter system) are located in the brain-suboesophageal ganglion complex, possibly in the brain itself.     

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.     

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

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

Repeated larval diapause and diapause-free development in geographic strains of the burnet moth Zygaena trifolii Esp. (Insecta,Lepidoptera)

Summary Zygaena trifolii is a long-day insect with temperature-dependent photoperiodic responses. All larval instars are sensitive to photoperiod; however, diapause may occur at the third larval stage or any subsequent larval instars. There were quantitative differences within populations in the threshold photoperiod for diapause induction. The diapause response was polymorphic, so that larvae might enter diapause at different instars under the same culture conditions. Furthermore, decreasing photoperiods below a critical daylength shifted the diapausing instar towards earlier stages. Geographic strains of Z. trifolii showed discontinuous clinal variation. Near the northern edge of the distribution [Cologne (Köln), FRG], there is first an obligatory diapause, mainly during early instars, and additional facultative (repeat) diapauses during later larval instars in subsequent years. In the southern part of its distribution, this burnet moth is partially bivoltine in the field with a facultative first developmental arrest and a decreased capacity for repeated diapause (Valencia, Spain; Marseille, France). Further experiments indicated that the photoperiodically controlled diapause reaction is also influenced by the number of photoperiodic cycles experienced during the period spent in each larval instar, which depends on temperature. The adaptive significance of obligatory and facultative repeated diapause, varying even among the offspring of a single female, may be to buffer the populations against the more extreme and, from year to year, unpredictable fluctuations in climatic conditions at the northern edge of the distribution.Abbreviations L₃ feeding 3rd larval instar - L₄D diapausing 4th larval instar - L₅D2 repeat-diapausing larval instar with second diapause at the 5th larval stage - LD light-dark cycle - KT shortday conditions (e.g. LD 8:16) - LT long-day conditions (e.g. LD 16:8)     

An active principle from the corpora cardiaca,corpora allata and suboesophageal ganglion of the locust,inducing egg diapause in Bombyx mori

The activity of the substance(s) which are contained in the cephalic endocrine organs of the locust which induce egg diapause in Bombyx mori was examined by implantation and injection of saline extracts of these organs. Extracts from the median and lateral neurosecretory parts of the locust brain were not effective in inducing egg diapause. Extracts of the corpora cardiaca, corpora allata, and suboesophageal ganglion of the locust induced diapause eggs in Bombyx pharate adults from which the suboesophageal ganglion had been removed. The first two extracts could induce egg diapause even in isolated abdomens of pharate adults of Bombyx. In the locust corpora cardiaca, the activity was present only in the glandular lobe and not in the nervous region. This activity decreased when the nervi corporis cardiaci I and II and of nervi corporis allati I were cut. Allatectomy also brought about a decrease in the activity in the glandular lobe which could not be restored by the injection of juvenile hormone. The activity in the corpora allata was enhanced slightly by the disconnection though not significantly.From these results, it is assumed that the corpora cardiaca, corpora allata and suboesophageal ganglion of the locust contain and active principle(s) capable of inducing egg diapause in Bombyx mori. The nervous connections between the brain, corpora cardiaca, and corpora allata are essential for the accumulation of the active substance(s) in the glandular lobes of the corpora cardiaca.     

The seasonal development cycle of <Emphasis Type="Italic">Aphantopus hyperantus</Emphasis> (L.) (Lepidoptera,Nymphalidae: Satyrinae) in Leningrad Province

The simultaneous influence of the photoperiodic and temperature conditions on pre-diapause and postdiapause larval development of the ringlet Aphantopus hyperantus was studied. At the short day (12 h of light a day) all larvae reached the III instar and entered diapause at all the temperatures tested (18–24°C). At the long day (22 h) the number of diapausing larvae increased with decreasing temperature, and the larvae diapaused at the III and IV instars. The I and II instar larvae (before winter) developed and grew faster under short-day, and the hibernated larvae, under long-day conditions. At the short day the growth rate of the I and II instar larvae did not depend on the temperature, and at the long day it decreased as the temperature increased. After hibernation the growth rate of the IV instar larvae increased with the temperature under long-day conditions, and did not depend on the temperature under short-day ones. Thus, acceleration or deceleration of development, depending on the photoperiodic and temperature conditions, regulated the timing of diapause in the overwintering stage (the III instar larva) and maintained the univoltine seasonal cycle. The males of A. hyperantus developed and grew faster than females while their weight at the larval and pupal stages was smaller. The thermal reaction norms for development (the lower temperature threshold and the coefficient of thermal sensitivity of development) were calculated for the eggs, IV and V instar larvae, and pupae. The coefficient of thermal sensitivity of development for the IV and V instar larvae taken together was lower, and the threshold was higher than the corresponding values for eggs and pupae. The thermal reaction norms for development of the latter stages were the same. The larvae of A. hyperantus retained their sensitivity to the photoperiod after hibernation. Resumption of development after cold reactivation occurred only under long-day conditions, whereas under short-day ones the overwintered larvae entered a repeated diapause. The possibility of photoperiodic reactivation and the absence of spontaneous resumption of development in these larvae were demonstrated.     

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.     

Alternative life cycles controlled by temperature and photoperiod in the oligophagous bug, Dybowskyia reticulata

Abstract. Effects of temperature and photoperiod on the induction and re-induction of adult diapause were examined in Dybowskyia reticulata (Dallas) (Heteroptera: Pentatomidae). Adults collected from the field after overwintering in early summer continued oviposition under long-day conditions of LD 16:8 h at 20 or 25°C, while they re-entered diapause under short-day conditions of LD 12:12 h at 25, 27.5 or 30°C. By contrast, adults reared in the laboratory from eggs at 20 or 25°C entered diapause under both long-day and short-day conditions, whereas those reared at 27.5 and 30°C entered diapause only under short-day conditions. Under quasi-natural conditions in 1993, when summer temperature was low, most adults of the first generation entered diapause in late July. However, in the warmer summer of 1996, oviposition was recorded in many females that ecdysed into adults from July to early August. Even though the seeds of the host plants occur in a restricted period from early summer to early autumn, in warmer years D. reticulata may produce a second generation. The response to temperature with a threshold between 25 and 27.5°C in D. reticulata brings about a switch between the univoltine and bivoltine life cycles.     

Photoperiodic induction in the silkworm, Bombyx mori, reared on artificial diet: Evidence for extraretinal photoreception

When a bivoltine race of the silkworm, Bombyx mori, was reared on artificial diets, the silkworm became photoperiodically responsive during larval development. The response curve for the embryonic-diapause induction of the next generation was of the long-day type, showing a sharply defined critical day-length of 16.5 hr. Complete blindness caused by covering the larval stemmata with black paint was confirmed electrophysiologically. It did not influence the diapause incidence. Vitamin A deficiency, which produced a similar defect in visual function in the stemmata, was also without effect on diapause. These observations show that extraretinal photoreception is involved in photoperiodic induction in the silkworm.     

Larval diapause in the mosquito aedes triseriatus: effects of diet and temperature on photoperiodic induction

Larval diapause was induced in both a northern (Ohio) and a southern (Alabama) strain of Aedes triseriatus by photoperiodic treatment of the larval stage itself. The effect of a short-day (10 hr) photoperiod during larval development was cumulative and resulted in failure of the fourth instar to pupate. The larvae, which had been obtained from long-day (16 hr) eggs, underwent diapause when their development was slowed by a low-quantity diet or low temperature (16°C) for a sufficient number of days to allow the short-day photoperiod to exert a visible effect.     

Role of the neuroendocrine complex in the control of adult diapause in the bean bug,Riptortus clavatus

In the bean bug, Riptortus clavatus, allatectomy suppressed reproduction in adults reared under nondiapause-inducing long-day conditions, and transection of the nervi corporis allati induced reproduction in adults reared under diapause-inducing short-day conditions. These effects of allatectomy and denervation were observed both in the morphology of reproductive organs and in the electrophoresis pattern of hemolymph proteins in both sexes. These results indicate that, in diapause adults, the brain suppresses the activity of the corpus allatum to secrete juvenile hormone through nervous pathways. The removal of the corpora cardiaca–corpus allatum complex in females not only inhibited ovarian development, as allatectomy did, but also prevented mature eggs in the oviduct from being laid. Therefore, it is assumed that the corpora cardiaca release an oviposition-stimulating substance. Arch. Insect Biochem. Physiol. 35:347–355, 1997.© 1997 Wiley-Liss, Inc.     

Photoperiodic response of larvae of the yellow-spotted longicorn beetle Psacothea hilaris after removal of the stemmata

The role of the stemmata in photoperiodism has been examined in holometabolic insects, but the only reliable results in Coleoptera have been obtained in Leptocarabus kumagaii (Carabidae), the larvae of which do not respond to photoperiod without stemmata. In the present study, photoperiodism was examined in another coleopteran, Psacothea hilaris (Pascoe) (Cerambycidae), after surgical removal of the stemmata. Larvae reared under short-day conditions and transferred to long-day conditions on day 2 of the 5th instar pupated without further larval molts, whereas those continuously reared under short-day conditions underwent supernumerary molts and did not pupate. When the stemmata were removed on day 2 of the 5th instar, the larvae pupated under long-day conditions but did not do so under short-day conditions. However, under long-day conditions some underwent supernumerary molts before pupation. Larvae from which the sensilla trichodeum were removed showed a similar response to that of stemmata-deficient larvae, and larvae from which stemmata were removed at a younger stage (day 2 of the 4th instar) responded to photoperiod similarly to intact larvae. Thus, supernumerary molts under long-day conditions after removal of the stemmata were attributed to injury due to surgery, rather than a change in photoperiodic photoreception. Therefore, we conclude that larvae of P. hilaris show a photoperiodic response after removal of stemmata, in contrast to larvae of L. kumagaii.