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.     

Effects of photoperiod and temperature on blood feeding,oögenesis and fat body development in the mosquito, Culiseta inornata

In the laboratory, blood feeding rates, oögenesis and fat body development were utilized to assess the effects of photoperiod and temperature in the conditioning of adult female Culiseta inornata for aestivation. Long-day (16L:8D) and short-day (8L:16D) photoperiods were examined in combination with temperatures of 15°C, 20°C, and 25°C. All tested individuals were reared from egg to adult under one of six possible regimes.Blood feeding occurred in approximately 60% of the females subjected to short-day conditions. Females subjected to long-day regimes exhibited gonotrophic concordance (the inhibition of blood feeding), evidenced by a decrease in overall blood-feeding rate to 20%. Inhibition of blood feeding by long-day females decreased in a linear fashion with increasing temperature. No interaction of the effects of photoperiod and temperature upon blood feeding was apparent. Examination of the ovarioles of post-blood feeding females, reared under each of the above conditions, revealed no evidence of gonotrophic dissociation (the inhibition of oögenesis despite continued blood feeding).Fat body hypertrophy occurred in females reared under long-day conditions, whereas hypotrophy of this tissue was apparent in short-day females. No significant difference in fat body development occurred between parous and nulliparous females reared under long-day photoperiod conditions. Short-day blood-fed females reared at 15°C deposited a significantly greater amount of fat than short-day blood fed females reared at 20° and 25°C, and short-day nonbloodfed females reared at 15°, 20°, and 25°C. The primary stimulus for fat body hypertrophy appears to be long-day photoperiod conditions. Temperature exerted little discernible effect upon this process, and there was no interaction between the effects of photoperiod and temperature upon the degree of fat body development.     

The regulation of development during diapause in Ephestia elutella (Hübner) by temperature and photoperiod

Diapausing larvae of Ephestia elutella reared at 20°C in short photoperiods (LD 11:13), and then maintained 12 weeks or longer at 5–15°C before transfer to 20 or 25°C, pupated sooner than unchilled controls. At 25°C, all samples kept in long photoperiods (LD 15:9) survived better and pupated faster than similarly treated samples held in short photoperiods (LD 9:15). Samples kept at 20°C after chilling pupated much slower than those at 25°C, and, except after exposure at 5°C, pupated at similar rates at LD 11:13 or 15:9, although mortality was higher at the shorter photoperiod. After exposure at 5°C, larvae required increased day-length as well as increased temperature to hasten pupation whereas after exposure at 10°C most responded to increased temperature only.For samples maintained in slightly heated or unheated outbuildings, the summer emergence was poorly synchronized and males on average emerged ahead of females. Samples moved from the unheated outbuilding to 25°C and long days in the laboratory in early spring, however, pupated quickly and males and females emerged together. A late phase of diapause development thus exists requiring both high temperature and long photoperiods to ensure a prompt resumption of morphogenesis. Spring temperatures in the United Kingdom are seldom high enough to synchronize the completion of diapause.     

Photoperiod influences reproduction in the prairie vole (Microtus ochrogaster)

Four experiments examined the role of photoperiod in the regulation of seasonal breeding in the prairie vole. Adult male voles maintained in short (8L:16D) as compared to long (16L:8D) photoperiods for 10 wk had reduced testicular and seminal vesicle weights, but fertility was not impaired. Male prairie voles reared from birth until 35 days of age in short as compared to long photoperiods also had reduced testicular and seminal vesicle weights, as well as diminished fertility. The incidence of pregnancy did not differ between long- and short-day female voles paired for 6 days with long- or short-day males (93%, 86%, 89%, and 88%, respectively). Photoperiod did not affect the incidence or the timing of postpartum pregnancies in long- or short-day females paired with long-day males through the birth of several litters. Adult male prairie voles retain only marginal sensitivity to short photoperiods, maturing males are highly responsive to short days, and adult females are insensitive to photoperiod. These data suggest that termination of the breeding season in the autumn may be due to the lack of fecund males in the population.     

Pupal diapause of Helicoverpa armigera (Lepidoptera: Noctuidae): sensitive stage for thermal induction in the Okayama (western Japan) population

Helicoverpa armigera (Hübner) exhibits a facultative pupal diapause, which depends on temperature and photoperiod. Pupal diapause is induced at 20 degrees C by short photoperiods and inhibited by long photoperiods during the larval stage. However, in some pupae (35% of males and 57% of females) of a non-selected field population from Okayama Prefecture (34.6 degrees N), diapause is not induced by short photoperiods. In the present experiment, the importance of temperature for diapause induction was studied in the non-diapausing strain, which was selected from such individuals reared at 20 degrees C under a short photoperiod of 10L:14D. Furthermore, the sensitive stage for thermal determination of pupal diapause was determined by transferring larvae of various instars and pupae between 20 degrees C and 15 degrees C. Diapause was induced by 15 degrees C without respect to photoperiod. When larvae or pupae reared from eggs at 20 degrees C under a short or a long photoperiod were transferred to 15 degrees C in the periods of the middle fifth instar to the first three days after pupation, the diapause induction rate was significantly reduced in both males and females, especially in females. In contrast, when larvae or pupae reared at 15 degrees C were transferred to 20 degrees C in the same periods, diapause was induced in males, but not in females. However, the diapause induction rate of pupae transferred to 20 degrees C on the fourth day after pupation was significantly increased in females. The results show that temperature is the major diapause cue in the photoperiod-insensitive strain and the periods of middle fifth larval instar to early pupal stage are the thermal sensitive stages for pupal diapause induction with some different responses to temperatures between males and females in H. armigera.     

Effects of photoperiod and temperature on the termination of diapause in the univoltine seed wasp Eurytoma plotnikovi

Abstract Mummified pistachios containing fully grown diapause larvae of Eurytoma plotnikovi Nikol'skaya (Hym., Eurytomidae) were collected in early August and late September in coastal northern Greece and subjected to various photoperiod and temperature treatments, then maintained at 19 or 26°C and a long-day (LD 16:8 h), a changing, or a short-day (LD 10:14 h) photoperiod until pupation. In larvae of early August (beginning of diapause) subjected for 20 weeks to 19°C under a long, a changing, or a short photophase, followed by 19°C and a long photophase, 50% of the larvae pupated after 24, 18 and 13 weeks respectively. After exposure for 20 or even 12 weeks to a short photophase and low temperatures (10 or 4°C), pupation occurred after only 7–8 weeks and was more synchronous. The ranges of temperature for diapause development and post-diapause morphogenesis overlap. After exposure for 12 weeks to short days and low temperature, larvae of late September pupated much sooner under long days than under short days and sooner at 26° than at 19°C. E.plotnikovi depends on both temperature and photoperiod for diapause development, low temperature having a strong favourable effect on the earlier part and long day on the later part of diapause. In a few larvae of another pistachio seed wasp, Megastigmus pistaciae Walker, after a long enough period of low temperatures, diapause was terminated normally at 26°C and long days, or at 19°C and long or short days.     

A method for rearing the yellow-spotted longicorn beetle, <Emphasis Type="Italic">Psacothea hilaris</Emphasis> (Coleoptera: Cerambycidae), to stabilize the last larval instar

The larvae of Psacothea hilaris (Pascoe) grown continuously under a long day in the laboratory pupate after the 4th, 5th, or 6th instar. This developmental polymorphism has complicated studies on the control of metamorphosis in P. hilaris. Since pupation in P. hilaris is known to be suppressed under a short day, a change in the photoperiod from a short to a long day with appropriate timing may assist in obtaining physiologically homogeneous larvae that pupate at the next molt. When the photoperiod was changed from 12-h light:12-h dark to 15-h light:9-h dark at the beginning of the 5th instar, the pupation rate at the next (5th) molt reached 90%, which was significantly higher than the proportion of larvae that pupated after the 5th instar when the larvae were grown continuously under a long day (52%). This rearing technique will expedite studies on the control of metamorphosis in P. hilaris.     

Effect of the photoperiod of the site of oviposition of Aedes mariae (Diptera,Culicidae)]

The overwintering biology of Aedes (Ochlerotatus) mariae (one of the sibling species of the mariae complex) was studied in populations from the Tyrrhenian coast of Central Italy (Sperlonga and Formia). Ae. mariae has in this zone several generations per year and its larvae are commonly found from March to October exclusively in rock pools along the coast. Field and laboratory observations show in these populations an embryonic winter diapause induced by short-day photoperiods (9-12 hours). The photoperiod acts primarily on the parental females during their preimaginal life. The expression of diapause is influenced by temperatures and photoperiods experienced by the embryos. Full embryonic diapause is observed when the eggs are incubated at relatively low temperatures (less than 16 degrees C) and at short photoperiods. Short-day photoperiods also induce a remarkable change in the oviposition behavior of Ae. mariae. Laboratory observations show that the adult females readily oviposit on water surfaces when originating from larvae reared at long-day photoperiod while they are very reclutant to oviposit in the same situation when reared at short day photoperiod. Choice experiments involving four alternative oviposition sites (see Plate 1) demonstrate a preference for outside free water in long-day mosquitoes and for inside moist surfaces in short-day mosquitoes. The behavioral difference persists in successive gonotrophic cycles and it is not apparently affected by the photoperiod acting on the adult females. The above evidence together with recovery of overwintering eggs in holes and crevices of rock pools suggest that the shift in oviposition site shown in the laboratory reflects a similar shift occurring in nature. Such photoperiodically induced change in oviposition behavior seems to have an important adaptive significance in providing more constant microclimates to the diapausing eggs and in protecting them from the mechanical action of winter storms.     

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.     

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.     

Effects of temperature on the development of overwintering immature stages of the near‐threatened butterfly Leptalina unicolor (Bremer & Grey) (Lepidoptera: Hesperiidae)

Overwintering larvae of multivoltine and univoltine populations of Leptalina unicolor were reared under various constant and fluctuating temperatures superimposed on a photoperiod of either 12 h of light and 12 h of darkness (LD 12:12) or LD 15:9. Diapause of the larvae terminated in midwinter (by early February). All the larvae of both populations pupated after two molts without feeding and the head capsule width of the final instar larvae was smaller than that of the penultimate instar ones. The photoperiod did not significantly affect larval development, but long‐day conditions (LD 15:9) hastened pupal development. The thermoperiod had a significant effect on the development of the multivoltine population. When multivoltine population larvae were kept under a low fluctuating temperature regime (cryophase/thermophase = 14/20°C), the period until adult eclosion was shorter than that under a constant temperature of 17°C. On the contrary, when larvae were kept under a high fluctuating temperature regime (24/30°C), the period until adult eclosion was longer than that under a constant temperature of 27°C. However, the univoltine population did not show such a reaction to the fluctuating temperature. The durations of final instar larva and pupa of the multivoltine population were shorter than those of the univoltine population. The developmental zeros of penultimate and final instar larvae and pupae of the univoltine population were lower than those of the multivoltine population. The head capsule width of penultimate instar larvae and the forewing length of adults of the univoltine population were larger than those of the multivoltine population for both sexes.     

A fine structural survey of the development of the adult fat body of Leptinotarsa decemlineata

The fat body of a female Colorado potato beetle, Leptinotarsa decemlineata, at adult ecdysis, contains a large number of protein granules which are composed of light and dark zones. Part of the light zone in some of these granules is believed to be urate. During the first two days after adult ecdysis, fat body development is not essentially different in females reared either under long- or short-day conditions. Protein granules and large vacuoles disappear and the first cell organelles are regenerated. The effect of the photoperiod on the histological structure of the fat is expressed after these events. In females reared under long-day conditions, the fat body becomes specialized for vitellogenin synthesis. Under short-day conditions, the fat body stores massive amounts of lipid until day 6 after adult ecdysis. Then the first electron-dense protein granules develop near the nucleus, and on day 10 the first autophagic vacuoles are seen. These structure changes are discussed in connection with the known biochemical properties of the adult faty body of Leptinotarsa.     

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.     

Influence of gonadal hormones on odours emitted by male meadow voles (Microtus pennsylvanicus).

Free-living male meadow voles (Microtus pennsylvanicus) emit odours that are attractive to females at the beginning, but not at the end, of the breeding season. The effect of gonadal hormones on female-attractant cues was examined in males born and reared in long (14 h light day-1) and short (10 h light day-1) photoperiods that simulate daylengths in the breeding and nonbreeding seasons, respectively. Gonadectomy affected the attractant properties of odours emitted by long photoperiod, but not short photoperiod, males. Long photoperiod females preferred odours of intact rather than those of gonadectomized long photoperiod males, and odours of gonadectomized long photoperiod males rather than those of intact short photoperiod males. Females did not show a preference between the odours of intact and castrated short photoperiod males. Gonadal hormone replacement in males affected female responses to the odours emitted by long photoperiod, but not short photoperiod, gonadectomized males. Long photoperiod females did not display a preference between odours of intact long photoperiod males and gonadectomized long photoperiod males treated with testosterone or oestradiol. We conclude that in spring and summer gonadal hormones increase attractiveness of male odours; this effect may require aromatization of testosterone to oestradiol. Substrates that control attractiveness of odour cues in male voles appear to be unresponsive to androgens during the nonbreeding season.     

Biological and biochemical differences between in vitro- and in vivo-reared Exorista larvarum

Quantitative and qualitative parameters of Exorista larvarum (L.) (Diptera: Tachinidae) reared on two insect-material-free artificial media and in the factitious host Galleria mellonella L. (Lepidoptera: Pyralidae) were compared. Significantly higher puparial yields and weights were obtained in both a milk-based and a veal homogenate-based medium than in the factitious host. Longevity and parasitization rates were not different between the in vitro- and in vivo-reared flies. Despite the greater puparial weight of the veal medium-reared E. larvarum females, the number of eggs laid by these females on host larvae was not higher than that of females reared under the other two rearing conditions. Moreover, in a complementary experiment, with homogeneous puparial weights of milk medium- and host-reared females, the former oviposited fewer eggs. Hence, puparial weight alone is not a reliable quality parameter for E. larvarum reared on artificial media. Lower amino acid content, with a deficiency in aromatic amino acids and an excess in proline, was found for in vitro third instar parasitoid larvae reared on both media compared to the in vivo-reared ones. These results suggest a correlation between the amino acid deficiency and imbalance of medium-reared larvae and the lower number of eggs laid by the females obtained.     

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.     

Mating and diapause in hibernating adults ofMenida scotti puton (Heteroptera: Pentatomidae)

InMenida scotti aduts which mate during hibernation, we examined the effect of photoperiod on diapause induction in terms of the developmental degree of reproductive organs and corpora allata. In a hibernating population, mating season and physiological changes related to dipause were investigated. When newly emerged adults were reared under long- or short-day conditions, reservoir of the ectodermal accessory gland and corpora allata in males developed, while the developments of ovary and corpora allata in females were suppressed under both conditions. This suggests that diapause of this species is induced only in females independently photoperiodic conditions during adult stage. In a hibernating population, mating was observed from early to late November and from late March to early May. Observations of the development of reproductive organs and corpora allata suggest that diapause is induced only in females in the former period, while both sexes are not in diapause in the latter period.     

Photoperiodic control of larval diapause in the yellow-spotted longicorn beetle, Psacothea hilaris: analysis by photoperiod manipulation

The photoperiodic control of diapause induction in the larvae of the yellow-spotted longicorn beetle, Psacothea hilaris (Pascoe), was investigated using a west Japan-type population collected from Ino, Kochi Prefecture, Japan. In this population, the larvae expressed a long-day photoperiodic response with a critical daylength between 13.5 and 14 h at 25 °C ; under a long daylength, the larvae pupated after the 4th or 5th instar, while the larvae entered diapause under a short daylength after 2.3 additional molts on average. When the photoperiod was changed from a short (L12:D12) to a long (L15:D9) daylength, pupation occurred in most of the individuals irrespective of the time of the change. When the photoperiod was changed from long to short at 1 or 2 weeks after hatching, all of the larvae entered diapause, whereas when the photoperiod was changed at 5 weeks after hatching or later, most of the larvae pupated. The 2 weeks exposures to a long daylength against a 'background' of a short daylength at various times revealed that the larvae of this insect are most sensitive to the photoperiod from 4 to 6 weeks after hatching.     

Sex-related adaptive responses of Populus cathayana to photoperiod transitions

Populus cathayana Rehd., a dioecious tree species, occupies a wide range of habitats in southwest China. Both males and females were grown under two regimes of photoperiod, from mid-length to short-day photoperiod (SD shift), or to long-day photoperiod (LD shift). SD shift triggered leaf senescence in both males and females by decreasing net photosynthesis rate ( A ), transpiration ( E ), and chlorophyll pigment ( Chl ), non-structural carbohydrate (NSC) and indoleacetic acid (IAA) contents, while increasing abscisic acid (ABA), malonaldehyde (MDA) and free proline (Pro) contents. The antioxidant enzyme (e.g. POD, CAT and SOD) activities and capability to maintain ultrastructural integrity also decreased under SD shift. Males exhibited faster leaf senescence than did females, as shown by greater decreases in A , E , Chl and IAA. However, males maintained a less senescent stage than did females, as indicated by higher values of A , Chl , NSC, IAA and antioxidant enzyme activities. Conversely, A , E , NSC and IAA contents and antioxidant enzyme activities were enhanced by lower O₂^•− in females, whereas reduced by higher O₂^•− in males under LD shift. Such sex-dependent responses of P. cathayana to photoperiod transitions showed that males and females possess different adaptabilities, which may relate to sex-specific leaf senescence speed under changing environments.     

Action of photoperiod at cold in diapausing Chrysopa carnea females (Chrysopidae,Neuroptera)

Summary To reveal the effect of photoperiod on insects under low temperatures, dormant females of Chrysopa carnea were exposed for 3 months (November–January) to cold (7°±2.5°C) and to long- or short-day conditions (18L:6D or 12L:12D). They were thereafter activated under short-day conditions at 25°C.The samples differed considerably in the incidence of ovipositing females, duration of pre- and post-oviposition period, and fecundity. All parameters evidenced a considerably higher reproductive activity of insects chilled under short-day conditions. It was thus confirmed that adults of C. carnea perceive photoperiod at cold. The action of short photoperiod at cold does not impair the subsequent oviposition; by contrast it promotes future reproduction, apparently also due to photoperiod changes. Although short-day photoperiod inhibits the activation at warm temperatures, it enables diapause development under cold temperatures.Dedicated to Prof. Dr. Hans Joachim Müller, Jena, on the occasion of his 65th birthday