Adult diapause ofCybocephalus nigriceps nigriceps [Col.: Cybocephalidae]

During autumn and winter (October–February)Cybocephalus nigriceps nigriceps (J. Sahlberg) adults undergo a facultative diapause. The diapausing adults agregate in hiding places, development of the ovaries is arrested, and prey consumption is lowered. Diapause can be prevented by exposure to a long-day photoperiod (16 hr light) at high temperatures (29° and 32°C). Diapause is induced mainly by short-day conditions, but the termination of diapause is affected by both photoperiod (long-day conditions) and temperature (29°, 32° and 34°C). In mature females, diapause induction causes ovarian degeneration. The induction or inhibition of adult diapause is affected by photoperiods occurring during larval development as well as during adult life.     

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.     

Effect of photoperiod and temperature on diapause in a Florida strain of the tropical warehouse moth Ephestia cautella

A photoperiodically-controlled diapause of the long-day, short-day type was identified in a brown-winged, yellow-eyed strain of Ephestia cautella (Walker). The proportion of larvae diapausing in very long photoperiods was less than in short photoperiods. The mean critical photoperiod, here defined as that photoperiod giving half the maximum percentage of insects that diapause in response to photoperiod at a given temperature, was between 12 and 13 hr for the long-day reaction at both 20 and 25°C. The principal sensitive phase occurred near the time of the last larval moult. The mean duration of diapause was 2–3 months at 20°C and slightly longer at 25°C. The optimum temperature for diapause development was near 15°C, all larvae pupating within 24 days after a 45-day exposure at this temperature. Diapause could be terminated whenever larvae diapausing at 20°C were exposed to as few as five long (15 hr) photoperiods at 25°C. Long photoperiods at 20°C, or short photoperiods (9 hr) at 25°C were less effective in terminating diapause.     

Photoperiodic and thermoperiodic interactions in the regulation of the larval diapause of Diatraea grandiosella

The effect of various combinations of photoperiod and temperature on the induction and termination of the mature larval diapause of a Missouri strain of the southwestern corn borer. Diatraea grandiosella, was examined. Larval exposure to regimes in which the low phase of a 30°:23°C thermoperiod coincided with a scotophase of 10 to 14 hr duration led to high incidence of diapause. Larval exposure to 30°:24°C, 33°:21°C, and 36°:18°C thermoperiods with half cycles of 12 hr in continuous darkness yielded a diapause incidence of 16%, 22%, and 59%, respectively, whereas exposure to a 30°:24°C thermoperiod in continuous illumination yielded a completely nondiapause generation. Larval exposure to one of a series of 36°:18°C thermoperiods in which the duration of the high phase was increased in 2 hr increments from 0 to 24 hr in continuous darkness showed that “short-day” thermoperiods yielded a high incidence of diapause. However, no clearly defined critical thermoperiod was observed. An examination of photoperiodic and thermoperiodic effects on diapause development showed that, in general, those combinations of temperature and light cycles which were diapause inductive also retarded diapause development. The relationship between seasonal photoperiods and thermoperiods in southeastern Missouri was examined.     

Aspects of the induction of diapause in a laboratory strain of the mite Tetranychus urticae

Some diapause characteristics were studied in a strain of the spider mite. Tetranychus urticae. which had been reared on bean plants in the laboratory for over 15 yr. The diapause induction response curve was of the long-day type, showing a sharply defined critical daylength of 13 hr 50 min. In constant darkness no diapause induction occurred, but with a photoperiod of 1L:23D diapause incidence was already complete. A thermoperiod with a 5°C amplitude induced diapause in combination with a short-day photoperiod only when the low phase of the thermoperiod coincided with the scotophase. The same thermoperiod did not induce any diapause in constant darkness. The photoperiodic reaction of the laboratory strain used in these experiments appeared to remain constant over a very long period of time and to be independent of the diapause history of previous generations of mites.Although photoperiodic sensitivity was demonstrated during the whole postembryonic development, sensitivity was maximal at the end of the protonymphal instar and declined rapidly during the deutonymphal instar. Only 2 inductive cycles of 10L:14D were required to induce up to 62% diapause if the mites were kept in continuous darkness during the remainder of their development. Long days or continuous light could reverse the inductive effect of a sequence of short-day cycles previously applied to the mites.Light breaks of 1 hr duration applied at different times during the dark period of a 10L:14D photoperiod generated a sharp bimodal response curve with two discrete points of sensitivity to the light breaks at 10 hr after ‘dusk’ and 10 hr before ‘dawn’, thus showing a remarkable similarity with the results obtained in light break experiments with some species of insects.     

Effect of photoperiod on a winter and on a summer diapause in two species of cranefly (Tipulidae)

Two craneflies, Tipula subnodicornis and Tipula pagana, both undergo diapause in the final larval instar. The species showed differences in the intensity of diapause and in the timing of the photoperiodic reaction during diapause, that could be related to season. Tipula subnodicornis undergoes a winter diapause that is induced and maintained in its early stages by short photoperiod (L:D:6:18). In the laboratory individuals in the early stages of diapause terminated diapause and pupated earlier when they were exposed to daylengths of, or greater than, 12 hr. However, it is suggested that in the field diapause is broken before the natural daylength is long enough to have any accelerating effect on development. Tipula pagana has a summer diapause which is of greater intensity than that of Tipula subnodicornis and some larvae were maintained for 197 days in the laboratory, without pupating, on an 18 hr daylength. Diapause was broken by a L:D;16:8 photoperiod and development was accelerated by a further decrease in daylength. The acceleration in development rate was attended by a decrease in the variance about the mean date of emergence and resulted in a highly synchronised emergence period. It is suggested that this quantitative response to daylength is particularly important to a species that emerges in the autumn when the temperature in the field is falling.     

The photoperiodic clock in the flesh-fly, Sarcophaga argyrostoma

Larval cultures of the flesh-fly, Sarcophaga argyrostoma, were raised in experimental light cycles with periods (T) of 21 to 72 hr, each cycle containing a photoperiod of 4 to 20 hr of white light. This ‘resonance’ technique revealed periodic maxima (～24 hr apart) of pupal diapause, thereby demonstrating an endogenous circadian component in the photoperiodic clock. The positions of these maxima of pupal diapause suggested that the oscillation, like that controlling the pupal eclosion rhythm in Drosophila pseudoobscura, is ‘damped out’ by photoperiods longer than about 11 to 12 hr, but restarts at dusk whereupon it runs with circadian periodicity in a protracted dark period. With photoperiods shorter than 12 hr, however, the two diapause maxima were less than 24 hr apart, suggesting that an additional component, possibly a ‘dawn hour-glass’, was modifying the position of the first peak.Both photoperiod and the period of the driving light cycle (T) were shown to affect the length of larval development (the sensitive period) and the number of calendar days needed to raise the incidence of pupal diapause to 50 per cent (the required day number, RDN). Peaks of diapause induction were shown to be the result of an interaction between a long sensitive period (slow development) and a low RDN, whereas troughs in diapause induction were the result of an interaction between a short sensitive period (fast development) and a higher RDN.Larvae of S. argyrostoma are unable to distinguish (in a photoperiodic sense) between 12 and 18 hr of red light (600 nm).     

Temperature and photoperiodic regulation of diapause of the southwestern corn borer,Diatraea grandiosella

The effect of temperature and photoperiod on the onset and termination of the mature larval diapause of the southwestern corn borer, Diatraea grandiosella, was examined. The results showed that diapause induction was an extremely temperature-dependent process. Larvae reared at 23°C under short days all entered diapause whereas 90 per cent of those reared at 27°C developed continuously. A photoperiodic response was only demonstrable at 25°C, when diapause was instituted following larval exposure to daily photophases ranging from 8 to 14 hr. An examination of the sensitivity of immature larvae to low temperatures revealed that all instars showed some susceptibility to low temperature induction of diapause. No intermediate instar was found in which the developmental programme could be switched entirely from a diapause to a non-diapause one, or vice versa.Diapause development was also found to be primarily temperature regulated. The rate of termination of diapause at 30°C was significantly higher than that at 25 or 23°C. Continuous exposure to light rather than a daily photoperiod produced the highest rate of diapause development. It was also shown that a period of chilling (5°C) did not accelerate diapause development. The significance of the results in relation to the evolution and geographical distribution of the species is discussed.     

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.     

Effect of photoperiod on the development and diapause of the green lacewing Chrysopa pallens (Neuroptera: Chrysopidae)

To investigate the physiology of Chrysopa pallens, the effect of photoperiod on diapause and development was examined in a Japanese population (33.4°N). The response stage for diapause of C. pallens was considered to be the prepupal stage. The critical photoperiod for diapause induction at 20.0°C was between 13 h light : 11 h dark (LD 13:11) and LD 14:10. The larval developmental period was affected by photoperiod: larvae in diapause took longer to complete their development. This difference of larval developmental period in relation to photoperiod was considered to be an adjustment of larval diapause timing.     

Control of summer and winter diapause in the leaf-mining fly Pegomyia bicolor Wiedemann (Dipt., Anthomyiidae)

Effects of photoperiod and temperature on diapause induction and termination were investigated in both aestival and hibernal pupae of Pegomyia bicolor Wiedemann under field and laboratory conditions. In the field, summer diapause had occurred already in part of the first pupal population; the proportion of diapause gradually rose as the day length and temperature increased. This fly is a short-day species with a pupal summer and winter diapause. Summer diapause was induced by both long day-lengths and mild temperatures. The whole larval life is sensitive to photoperiod. Winter diapause was induced mainly by low temperatures, especially in the first 10 days after pupation. High temperatures strongly enhanced summer diapause induction regardless of photoperiod. The diapause-averting influence of short photoperiods was fully expressed only at moderately low temperatures. High temperatures delayed diapause development, resulting in a rather long summer diapause; whereas low temperatures hastened it, leading to a short winter diapause and showing a low thermal threshold for diapause development. In the field, the post-diapause development started in January, the coldest month, suggesting that the thermal requirements for post-diapause development is also low.     

Why is the number of days required for induction of adult diapause in the linden bug Pyrrhocoris apterus fewer in the larval than in the adult stage?

Adult females of Pyrrhocoris apterus, programmed for diapause by short-day (SD) photoperiod and those programmed for reproduction by long-day (LD) retain photoperiodic information in continuous darkness (DD) until death. However, if the interruption of SD by DD is made in the course of diapause programming in adults, then the incidence of diapause depends on the number of SD cycles received before DD, with no evidence that the photoperiodic clock is free-running at DD to complete diapause induction. These results indicate that the photoperiodic clock is stopped after transfer to DD and the information accumulated before transfer to DD is maintained. Diapause programming in the adult stage requires 9–10 SD cycles to induce diapause in 80% of individuals. However, if the diapause programming starts after ecdysis of LD-larvae to the last instar, only 3 SD cycles before transfer to DD are required for diapause in 80% of individuals. Surprisingly, if the newly ecdysed last instar LD-larvae, sensitive to photoperiod, are transferred to DD (thus they did not experience any SD), diapause occurs in 40% of the individuals. Thus, diapause ‘information’ is present in LD-larvae and is responsible for a lower number of SD required for diapause induction in the larval than in the adult stage.     

Critical photoperiod and daylength threshold differences between northern and southern populations of the butterfly Limenitis archippus

Laboratory studies show that different photoperiods induce diapause in northern (Vermont) and southern (Maryland) larval strains of the butterfly Limenitis archippus. The northern strain responds to $\text{1}\text{2}\text{hr}$ longer photoperiod thresholds and critical ranges than does the southern one. These responses are correlated with geographic differences in the ambient photoperiod of the two localities. In this facultative diapausing species, third instar larvae construct hibernacula within the basal portions of tubular leaves spun with silk, when daylength approaches either 13·5 hr (Vermont strain) or 13·0 hr (Maryland strain). When reared in total darkness some larvae develop directly to fourth inszar without diapause, although mortality is high. Among both strains different broods exhibit different incidences of diapause. Reciprocal inter-strain hybrids show intermediate diapause responses, suggesting that larval diapause is under the control of multiple genes.     

Diapause induction in the oblique-banded leafroller Choristoneura rosaceana (Lepidoptera: Tortricidae): Role of photoperiod and temperature

Induction of diapause in the larval stage of the oblique-banded leafroller, Choristoneura rosaceana (Harris), was found to be dependent on both photoperiod and temperature. At constant temperatures of 24, 20 and 16°C, short photoperiods induced diapause. The critical photoperiod was between 14–15 h of light per day at 20 and 16°C. At 14 h light: 10 h dark, all larvae expressed diapause. Temperature had a modifying effect, and slightly shifted the larval response to diapause-inducing photoperiods. High constant temperatures of 28°C and above induced diapause in some individuals (< 20%), while fluctuating temperatures of 32 and 16°C in a 12-h cycle resulted in 67% diapause induction, suggesting that diapause could also be induced by fluctuating temperatures, particularly if the higher temperature exceeds 25°C.The first- and the second-instar larvae were the only two stages sensitive to diapause induction. Exposure of adult, egg and third, fourth, and fifth-larval instars to diapause-inducing conditions did not produce diapause. Although diapause was induced in the first or the second instars, it was always expressed in the third or fourth instar.     

Effect of photoperiod and temperature on the induction of diapause in a Japanese strain of <Emphasis Type="Italic">Aphidoletes aphidimyza</Emphasis> (Diptera: Cecidomyiidae)

The aphidophagous gall midge Aphidoletes aphidimyza (Rondani) (Diptera: Cecidomyiidae), a dominant natural enemy of aphids, has been used as a biological control agent in many countries to control aphids in greenhouses. As developmental arrest in diapause lowers the effectiveness of natural enemies, we studied the effect of photoperiod and temperature on the incidence of diapause in a Japanese strain of A. aphidimyza by examining diapause induction under different day-length conditions in the laboratory. The critical day length for diapause induction was determined to be 12.7 h at 20°C. Diapause incidence was completely prevented at 30°C even though the photoperiod used was 11L13D. We also examined diapause induction under changing temperature conditions while maintaining the critical day length (12.7L11.3D). Diapause incidence was 100% in both field and greenhouse conditions under alternating temperatures of 20/16 or 25/16°C while the critical day length of 12.7 h was maintained. The Japanese strain of A. aphidimyza was sensitive to diapause entry from the first to the third noncocooned instar larval stages. Its eggs do not seem to be sensitive to diapause induction. Our results suggest that constant short-day conditions for at least four days are needed to induce diapause in the Japanese strain of A. aphidimyza.     

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.     

Seasonal plasticity in growth and development of the speckled wood butterfly, Pararge aegeria (Satyrinae)

Regulation of growth and development by photoperiod was studied in a population of the speckled wood butterfly, Purarge aegeria L. (Lepidoptera: Satyrinae), from southern Sweden. Individuals were reared in a range of photoperiodic regimes (9L. to 22L) and temperatures (13°C to 21° C). Plasticity was found for important life-history traits- generation time, growth rate and final weight and seasonal regulation of development in response to photoperiod was found to occur at two levels. Purarge aegeria hibernates as a third instar larva or in the pupal stage, cantering one of four major developmental pathways in response to photoperiod: (1) direct development in both the larval and pupal stages, (2) pupal winter diapause with or (3) without a preceding larval summer diapause, or (4) larval winter diapause. In addition to this high-level regulation of individual development, larval growth rate and pupal development rate also appear to be finally regulated by photoperiod within each major pathway. As photoperiods decreased from 22 h to 17 h at 17° C, growth rate among directly developing larvae increased progressively, as was the case for larva? developing according to a univoltine life cycle from 17 h to 14 h. At two photoperiods, 13 h and 16 h (corresponding to shifts between major pathways), both larval and pupal development were extremely variable with the fastest individuals developing directly and the slowest developing with a diapause. This indicates a gradual nature of diapause itself, suggesting that the two level may not he fundamentally different.     

Induction and development of winter larval diapause in a drosophilid fly,Chymomyza costata

Photoperiodic response during induction of larval hibernal diapause of Chymomyza costata was characterized and the course of diapause development was analyzed in the laboratory. C. costata becomes sensitive to photoperiodic stimuli during an unspecified stage of its early development (embryo, 1st larval instar); the sensitivity gradually increases during the 2nd and early 3rd larval instars and reaches its maximum just before the moment when it abruptly ceases at the age of 15-19 days after oviposition. Diapause intensifies during a period of 2-3 weeks after induction and, later, is maintained without apparent development until death (between 150 and 250 days) under 18 degrees C and a short-day photoperiod (L10:D14, SD). Diapause may be terminated in a horotelic process by exposure to a low temperature (2 degrees C) during which larvae subsequently (1) synchronize their post-diapause development (requires up to 14 days of chilling), (2) lose photoperiodic sensitivity (2 months), and finally (3) terminate diapause (5 months). Alternatively, diapause may be terminated in a tachytelic process by exposure to a high temperature (18 degrees C) and long-day photoperiod (L16:D8, LD) during which no synchronization occurs and pupariation takes place after a mean of 25.2 days (with a broad range from 8 to more than 50 days). Larvae that are transferred from LD to SD during their sensitive period switch their developmental programming from pupariation to diapause. Proliferation of adult primordial structures (imaginal discs, neuroblasts) slows down within 1 day after transfer. In contrast, whole body growth continues for at least 3 days before its rate slows down and matches the rate characteristic for SD conditions.     

Effect of temperature on photoperiodic response in a selected 'non-diapause' strain of Pyrrhocoris apterus (Heteroptera)

Abstract. The artificially selected 'non-diapause' strain of Pyrrhocoris apterus (L.) (Heteroptera) showed no diapause response to photoperiod at 26°C (Socha & Hodkova, 1994). However, the diapause response to short-day photoperiod (LD 12:12 h) became apparent at lower temperatures of 17°C (70% diapause) or 20°C (41% diapause). Diapause was induced in 60% females by short-day photoperiod combined with thermoperiod of 26/16°C, whereas only 20% diapause was induced by the same thermoperiod under continuous darkness. Thus the time-measuring system was not removed by artificial selection but the diapause response was shifted to lower temperatures. The diapause response to short days seems to be favoured rather by low temperature during scotophase than by low temperature throughout the whole light/dark cycle. If the percentage of diapause at 26°C is compared in F₁ hybrids and in wild and selected parental strains the diapause appears to be dominant at LD 13:11 h but recessive at LD 11:13 h and LD 10:14 h. A hypothesis is proposed that the inheritance of the percentage of diapause in F1 hybrids is determined by interactions of genes controlling the temperature dependence of photoperiodic response.