Effect of changes in the thermoperiod on reproductive diapause in Bruchidius atrolineatus Pic (Coleoptera: Bruchidae)

Abstract. Bruchidius atrolineatus (Pic) is a tropical bruchid developing in Vigna unguiculata (Walp) pods. Adults are in reproductive diapause during the dry season. In conditions of 40:25C or 35:25C, diapause is induced when larvae develop in thermophases longer than 14 h, in thermophases shorter than 10 h and in continuous darkness. Thermophases of 12 h cause a reduction of the proportion of diapausing beetles. This proportion also depends on the thermoperiod temperatures. Development at low temperatures (23: 16C, 12: 12 h) induces a high percentage of diapausing beetles. However, at low temperatures, the developmental times of the beetles are long and show important interindividual variability. When larvae develop in decreasing temperatures, small changes in thermoperiod temperatures (35:25C to 3O:2OC) have a strong effect on diapause induction. When larvae develop at low temperatures (25:15C) and then at high temperatures (40:25C), the proportion of diapausing beetles depends on the duration of development at low temperatures. Whatever the developmental conditions, both sexually active and diapausing beetles always emerge from the seeds. This interindividual variability is important in this species which has a wide geographical distribution in Africa.     

Induction of reproductive diapause in Bruchidius atrolineatus during the dry season in a Sahelian zone

Bruchidius atrolineatus (Pic) (Bruchidae: Coleoptera) is a bruchid beetle infesting Vigna unguiculata (Walp). During the dry season, adults are in reproductive diapause. Studies of adult populations in a traditional store containing contaminated pods of V. unguiculata show that the reproductive diapause is induced during the first larval instars. The temperature decrease from November to January probably is the main climatic factor inducing this diapause. The percentage of diapausing adults depends on the duration of the larval development at low temperatures. Experiments in climatic boxes confirm these results. At 12 h 25 °T-12 h 20° C 11–13 LD, a high proportion of the adults emerging from the V. unguiculata seeds are in reproductive diapause and the duration of their growth is variable. L1 L2 L3 larvae probably are sensitive stages to the thermal variations and the processes allowing emergence of diapausing adults probably are induced during these larval instars.

---

Résumé Bruchidius atrolineatus (Pic) est un Coléoptère Bruchidae se développant en zone sahélienne aux dépens d'une légumineuse Vigna unguiculata. Pendant la saison sèche, les adultes sont en diapause reproductrice. Un suivi des populations d'adultes a été réalisé dans un grenier traditionnel en terre séchée contenant des gousses récoltées dans la nature et portant des ufs de B. atrolineatus. Il a permis de constater que la diapause reproductrice était induite durant les premières phases du développement larvaire. La décroissance des températures de Novembre à Janvier avec des minima de 15 ° C et des maxima de 25 °C est probablement le principal facteur permettant l'induction de cette diapause. L'introduction dans le grenier de gousses portant des ufs venant d'être émis à différentes périodes de la saison sèche montre que le pourcentage d'adultes diapausants dépend de la durée du développement larvaire aux températures basses. Des expériences réalisées dans des étuves programmées confirment ces résultats. A 12 h 25 °T/12 h 20 °C 11–13 LD, un taux élevé d'adultes émergeant des graines est en diapause reproductrice et la durée de leur développement post-embryonnaire est très variable. En conditions plus chaudes 12 h 40 °C/12 h 25 °C 11–13 LD, le taux d'adultes diapausants est plus réduit et la durée du développement plus courte. L'analyse de la croissance larvaire montre que les premiers stades larvaires sont probablement des stades sensibles aux variations thermiques. Les modifications physiologiques permettant l'émergence d'adultes diapausants ou sexuellement actifs seraient induites durant ces stades.

     

A female sex pheromone in Bruchidius atrolineatus (Pic) (Coleoptera, Bruchidae): analysis of the factors affecting production or emission of this pheromone

ABSTRACT. . The females of the tropical Bruchid Bruchidius atrolineatus (Pic) (Coleoptera) produce a sex pheromone that influences male behaviour. This pheromone, tested on males in an olfactometer, triggers characteristic antennal movements followed by a distinct chemo-anemotaxis. Sex pheromone production depends on the age of the females and is partly correlated with their ovarian activity. Females 0–24 h old are not attractive, but they begin to attract males on the second day of adult life. Vitellogenesis seems to correlate with the emission or the production of the pheromone but there is no precise synchrony between the two phenomena which are probably induced by the same endocrine factors. Within 1 h after mating there is a decrease in the females' attractiveness, which lasts at least 48 h. Under our experimental conditions no particular rhythm of production or emission of the sex pheromone in B. atrolineatus could be observed.     

Effect of larval density on development and induction of reproductive diapause in Bruchidius atrolineatus

In Bruschidius atrolineatus (Pic), an increase in larval density inside the seed led to a reduction in the survival rate during post-embryonic development, with only a limited decrease in the weight of adults. A high larval density increased the proportion of adults in reproductive diapause under certain thermoperiodic conditions. Under other conditions that already promote the emergence of a high proportion of diapausing beetles, an increase in larval density had no effect on diapause.There was no correlation between the larval developmental time and the weights of adults in a strain with a very low incidence of diapause. However, such a correlation was observed when both sexuallyactive and diapausing beetles emerged. This correlation was explained by the longer developmental times and lower weights of diapausing beetles.

---

Résumé Chez Bruchidius atrolineatus, l'accroissement de la densité larvaire à l'intérieur de la graine entraîne une augmentation de la mortalité alors que la réduction de poids des individus à l'émergence est relativement limitée. L'accroissement de la densité larvaire provoque une augmentation du taux d'adultes en diapause reproductrice, dans certaines conditions thermopériodiques. Dans d'autres conditions, permenttant l'émergence d'un fort taux d'adultes diapausants, l'effet de la densité larvaire est nul. Lorsque l'on analyse l'effet de la densité chez les individus d'une souche ne présentant pas de phénomène de diapause, il n'y a pas de corrélation entre la durée de développement et le poids des adultes. Par contre, lorsqu'il y a émergence d'adultes sexuellement actifs et diapausants, il existe une corrélation entre la durée de développement et le poids des adultes émergeants. Cette corrélation est due au fait que les adultes diapausants, qui présentent les durées de développement les plus longues, sont ceux qui ont les poids les plus faibles.

     

Effects of photoperiod, temperature, and host plant age on induction of reproductive diapause and development time in Danaus plexippus

Abstract 1. Diapause induction in monarch butterflies Danaus plexippus was studied using adults captured from the wild in Minnesota and Wisconsin, and individuals reared under outdoor and controlled conditions.
2. Oocyte presence in females and ejaculatory duct mass in males were used to indicate reproductive status.
3. Some wild individuals were in diapause in mid-August, and all males and females were in diapause by late August and early September respectively.
4. Individuals reared under decreasing day lengths and fluctuating temperatures were more likely to be in diapause than were individuals reared under long or short day lengths or constant temperatures.
5. Individuals fed potted old Asclepias curassavica plants were more likely to be in diapause than were those fed potted young host plants; when cuttings of Asclepias syriaca plants from the field or greenhouse pots were used, there was no effect of host plant age.
6. Extremely high temperatures increased the number of day-degrees required for development from egg to adult, while decreasing day lengths and older host plants tended to decrease the number of day-degrees required for development.
7. There appears to be a continuum of reproductive development in monarchs, with gradual declines in mean ejaculatory duct mass and oocyte production during the late summer.
8. None of the experimental treatments led to 100% diapause, and diapause was more likely to occur in monarchs subjected to more than one diapause-inducing cue.     

Rhythmicity of mating and oviposition inCallosobruchus subinnotatus (Pic) (Coleoptera: Bruchidae)

Mating and oviposition behaviors were studied inCallosobruchus subinnotatus. Copulation was most frequent during the late scotophase, 2–3 h before onset of photophase. The females were less willing to mate during photophase, which increased the time to initiate mating while decreasing the duration of mating. Females exhibited increased movement prior to mating, resting immediately after mating, and remained stationary for 6 h when oviposition commenced. Multiple mating by both males and females affected the number of eggs laid, duration of mating, and uncoupling time at the end of mating. Females that mated two or three times laid more eggs than females that mated once or more than three times. Females that remainedin copula for less than 18 min showed greater readiness to remate than those that remainedin copula longer. There was a gradual decrease in the number of eggs females could lay with an increase in the number of previous matings by males.     

Effects of thermoperiod on photoperiodic determination of larval diapause in Ostrinia nubilalis

Some effects of thermoperiods on the photoperiodic determination of larval diapause in the European corn borer, Ostrinia nubilalis are reported. Thermoperiods and photoperiods were shown to interact to a highly significant degree in the induction of diapause. Diapause determination was found to be sensitive to the duration and temperature of the cryoscotophase of the thermophotoperiod; the thermophotophase characteristics exerted only minor influence. Cryoscotophase temperatures of 10°C or lower caused an unexpected increase in the apparent critical nightlength for diapause determination. A 0°C cryoscotophase effectively prevented the photoperiodic determination of diapause. Experimental investigation of the effects of 0°C cryoscotophases produced evidence that such low-temperature cryoscotophases may suppress the biological clock functions regulating the determination of diapause. Photoperiodic regimes in which 0°C pulses were inserted at successive points in 24-h cycle evoked responses suggesting that the early scotophase (saturation phase) was more strongly influenced by the low temperature pulse than was the later scotophase (scotonon); low temperature had little effect on the light-dependent processes (photonon) associated with the photophase.     

Effect of photoperiod and thermoperiod on the eclosion rhythm ofTrichogramma evanescens

The dynamics of adult eclosion inTrichogramma evanescens Westw. were studied under (1) constant light and temperature of 20 °C, (2) photoperiods L12:D12 and L16:D8 at 20 °C, (3) thermoperiod 4 h 27 °/20 h 20 °C at constant light, (4) joint action of photo- and thermoperiod. The emergence was arhythmic in constant light combined with constant temperature, while a sharp monophasic rhythm was observed with the isolated action of photo- or thermoperiod. The ‘light-on’ and ‘temperature step-up’ signals were shown to act in one direction. When both signals were combined, they manifested themselveses competing entraining stimuli which, in turn, revealed an apparent individual variation in relative reactivity to the light and temperature signals. Some perspectives which follow from these observations are discussed.     

Influence des inflorescences de la plante hôte (Vigna unguiculata) sur la levée de la diapause reproductrice de Bruchidius atrolineatus

Bruchidius atrolineatus (Pic) présente en zone sahélienne une diapause reproductrice durant la saison sèche et une partie de la saison des pluies. Les femelles diapausantes ne produisent pas de vitellogénine et le germarium des ovarioles est seul développé. Chez les mâles la spermatogenèse est très ralentie et les glandes annexes sont inactives. Lorsque les insectes diapausants sont placés en présence d'inflorescences de Vigna unguiculata Walp, leurs organes reproducteurs deviennent fonctionnels après un temps de latence de 15 à 20 jours. Il n'y a par contre aucune levée de la diapause chez des bruches placées en présence de gousses sèches de V. unguiculata dans une atmosphère saturée. Des informations sensorielles issues des pièces florales ou (et) des facteurs nutritionnels sans doute liés a la consommation de nectar semblent être à l'origine de cette levée de la diapause. Le pollen fort peu consommé n'a par contre aucun rôle. Cette régulation du cycle reproducteur de B. atrolineatus par les inflorescences de V. unguiculata permet l'émission des oeufs dès que les gousses commencent à se former à la fin de la saison des pluies.

---

Summary Bruchidius atrolineatus (Pic) is a widely distributed bruchid in the Sahelian zone which shows a reproductive diapause during the dry season and part of the rainy season. Diapausing females do not produce vitellogenin and their ovaries are reduced to the germarium. Spermatogenesis is very much reduced and male accessory glands are inactive. When these insects were placed in the presence of inflorescences of Vigna unguiculata which were renewed daily, the reproductive diapause of both males and females was interrupted after 15–20 days. Vitellogenesis occurred in the females and spermatogenesis increased in the males whilst their accessory glands became functional. When diapausing bruchids, found in stores of on V. unguiculata seeds during the dry season, were placed near the host plant's inflorescences, diapause was also terminated. In all cases, diapause was not interrupted when the insects were offered dry pods of V. unguiculata in a water-saturated atmosphere. The pollen, which is hardly eaten by this bruchid, did not seem to stimulate termination of diapause. Sensory stimulations derived from the flowers or/and nutritional factors may be the cause of the development of the reproductive organs. After termination of the diapause the males showed normal sexual activity whereas female fecundity was rather low, at least in our experimental conditions. This type of reproductive regulation allows this sahelian bruchid to resume its sexual activity when the host plant's flowers appear in the field at the end of the rainy season. Then the beetles lay their eggs on the pods as soon as the pods are developed.

     

The effect of cold temperature exposure and long-day photoperiod on the termination of the reproductive diapause of newly emerged female Pissodes strobi (Coleoptera: Curculionidae)

Abstract

• 1 There is confusion in the literature concerning a possible reproductive diapause in the adult white pine weevil Pissodes strobi.
• 2 We evaluated the effects of temperature, photoperiod, feeding substrate and mating status on the sexual maturation and oviposition of female white pine weevils.
• 3 Less than 30% of female P. strobi became sexually mature and laid eggs without experiencing dormancy under a temperature regime of 2 °C for 4 weeks.
• 4 Among the females that experienced a cold temperature treatment after emergence, 80% laid eggs after dormancy when exposed to a long‐day (LD 16 : 8 h) photoperiod and 17.6% laid eggs when exposed to a short‐day (LD 8 : 16 h) photoperiod.
• 5 Significantly more eggs were laid by all the females (with and without a cold treatment) when subjected to a long‐day photoperiod compared with a short‐day photoperiod.
• 6 A period of cold temperature followed by exposure to a long‐day photoperiod with warmer temperatures is required to break reproductive diapause and to obtain a good oviposition response in female P. strobi.
• 7 This study reveals the existence of much intraspecific variation in the response of the white pine weevil to temperature and photoperiod with respect to the induction and termination of reproductive diapause.

     

Analysis of multiparasitism by Eupelmus vuilleti (Craw) (Eupelmidae) and Dinarmus basalis (Rond) (Pteromalidae) in the presence of one of their common hosts,Bruchidius atrolineatus (Pic) (Coleoptera Bruchidae)

Bruchidius atrolineatus (Pic) is a tropical beetle (Coleoptera Bruchidae) that develops during the larval and pupal stages in the seeds of a legume Vigna unguiculata (Walp). Two species of Hymenoptera, Dinarmus basalis (Rond) and Eupelmus vuilleti (Craw), solitary ectoparasitoids of the larvae and pupae of B. atrolineatus, were introduced successively in the presence of their hosts, varying the interval between the two introductions. When D. basalis females were introduced 24 h, 3 days or 7 days after E. vuilleti, multiparasitism was low. The females had low fecundity, and their eggs were not distributed randomly over the different available hosts. When E. vuilleti females were introduced second, they oviposited on the different hosts availabe and did not avoid multiparasitism. The presence of hosts already parasitised by D. basalis increased the reproduction of E. vuilleti, and the fecundity of the females was higher than in control batches with E. vuilleti alone. E. vuilleti seems capable of detecting the ovipositor shafts drilled by the D. basalis females, and by introducing its own ovipositors killing the D. basalis eggs or larvae. When interspecific competition was occurring the number of E. vuilleti adults emerging from the seeds was no different from that observed in control batches with E. vuilleti alone, and there were always fewer D. basalis adults than in control batches (D. basalis alone). This interspecific competition reduces the influence of the two parasitoids in the biological control of bruchid populations.     

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.     

Effect of day length on development and reproductive diapause in Nysius huttoni White (Heteroptera, Lygaeidae)

Abstract:  Nysius huttoni White is endemic to New Zealand and an important pest of wheat and brassica crops. To provide critical information for pest forecast, management and quarantine inspection, we investigated the effect of day length on the growth, development and reproductive diapause of this pest under a series of photoperiodic regimes: 16 : 8, 14 : 10, 12 : 12 and 10 : 14 h [light : dark (L : D)]. Long day lengths [16 : 8 and 14 : 10 h (L : D)] promote a continuous lifecycle while short day lengths [12 : 12 and 10 : 14 h (L : D)] slow up the growth and development, prolong the pre-mating period, and induce the reproductive diapause. The absence of oviposition for approximately 30 days is recognized as the criterion for N. huttoni reproductive diapause definition. When all life stages are kept under the short day length conditions, only 60–73% of females enter reproductive diapause; if the exposure to short day lengths starts in late instar nymphs, 100% of females enter reproductive diapause. If only adults experience short day lengths, does diapause incidence fall in between, with up to 26% of females laying a few eggs before entering diapause. The critical photoperiod for the induction of reproductive diapause falls between 13.3 : 10.7 and 13.5 : 10.5 h.     

The effect of larval temperature and photoperiod on the incidence of larval diapause in the blowfly, Calliphora vicina

ABSTRACT. The blowfly, Calliphora vicina Meigen (Diptera: Calliphoridae), displays a maternally induced larval diapause. Progeny of adults exposed to short days enter diapause if the larval temperature is 15C or below; exposing adults to long days, or larvae to temperatures above 15C, results in non-diapause. By keeping progeny from short-day parents at the parental temperature of 23.5C for various lengths of time before transfer to 11C, it could be shown that the diapause-averting effect of high temperature operates in the late wandering stage of larva, possibly at the moment of the decision for the brain-ring gland complex to release PTTH/ecdysone at the diapause/non-diapause stage.
Although photoperiodic induction in the blowfly is maternal, previous work had shown that the larvae were also slightly sensitive to photoperiod. From the present study it is evident that also the eggs are photo-sensitive.     

The critical field photoperiod inducing ovarian diapause in the cabbage whitefly, Aleyrodes proletella (Homoptera: Aleyrodidae)

ABSTRACT. The ovarian and fat body development of cabbage whitefly, Aleyrodes proletella L., has been determined from field samples taken during the transition from a summer to an overwintering population. Between mid and late September 1983, the state of ovarian development decreases as the proportion of diapausing females increases. This is accompanied by maximum fat body development. Diapausing females emerge from late August onwards, and the critical photoperiod, based upon known larval sensitivity and including civil twilight as part of the photophase, is LD 15.5:8.5h. This is in agreement with laboratory studies. The majority of overwintering females are mated. Very few males survive the autumn.     

A genetic analysis of photoperiodic reproductive diapause in Drosophila triauraria

Abstract. Recombinant inbred lines were produced from hybrids between diapausing and non-diapausing strains of Drosophila triauraria Bock & Wheeler by twenty generations of full-sib mating. The analysis of these inbred lines suggests that the difference in the photoperiodic response between the original diapausing and non-diapausing strains is due to genes at three or four loci and at least one of these loci is located on the X chromosome. The original diapausing strain and two inbred lines exhibited circadian peaks when subjected to the Nanda-Hamner protocol, although they had different critical daylengths. This suggests that critical daylength is modified independently of the circadian oscillation system in this species. On the other hand, diapause duration was correlated with critical daylength among the inbred lines and the original diapausing strain, suggesting that these two traits are under the control of the same genetic system.     

Geographic variation of reproductive diapause in the Drosophila auraria complex (Diptera: Drosophilidae)

ABSTRACT. In the Drosophila auraria complex (D. auraria Peng, D. biauraria Bock & Wheeler, D. triauraria Bock & Wheeler, D. quadraria Bock & Wheeler and D. subauraria Kimura), D. triauraria from the island of Yaku (30.3N in latitude) and D. quadraria from Taiwan (24.1N) had no or weak diapause, while strains of all the four species from the main islands of Japan had a firm reproductive diapause. These strains entered diapause at short daylengths and the critical daylength was longer and the incidence of diapause higher at a low rather than high temperature. In each species, the critical daylength was longer in strains from higher latitudes. Among sympatric populations of different species, D. subauraria always had the longest critical daylength, while D. auraria had the shortest one. The populations of D. biauraria and D. triauraria had longer critical daylengths than the sympatric populations of D. auraria , but this difference became smaller northwards. Since a longer critical daylength induces diapause earlier in autumn, D. subauraria would enter diapause earliest among these species, and D. auraria would do so latest. The late start of diapause in D. auraria may be related to its adaptation to domestic environments. On the other hand, the adaptive significance of the late start of diapause in northern populations of D. biauraria and D. triauraria is not clear.     

Aestivo-hibernal reproductive diapause in Drosophila moriwakii: flight as a controlling factor

Drosophila moriwakii Okada & Kurokawa (Diptera; Drosophilidae) generally has only one generation per year and enters aestivo-hibernal reproductive diapause in Sapporo, northern Japan, but a small fraction of the population produced a second generation in summer at a place where breeding resources were abundant. In this species, diapause seems to be controlled by flight activity. When flies were cultured in cages in which they were able to fly freely, they entered diapause irrespective of photoperiod, but they did not do so at long daylengths when cultured in small vials in which they were prevented from flying. Furthermore, flies with wings removed did not enter diapause at long daylengths even if they were cultured in the cages.     

Influence of photoperiod on low temperature acclimation for cold-hardiness in Drosophila auraria

ABSTRACT. Imagines of Drosophila auraria Peng, a reproductive diapause species, developed cold-hardiness at low temperatures to a greater extent when exposed to a diapause-inducing photoperiod (LD10:14 h) than when exposed to a diapause-preventing photoperiod (LD 16:8h). Imagines kept at 18°C, which was the temperature at which they were reared to eclosion, did not survive a test exposure to -5°C for 8 days regardless of age or photoperiod. When transferred to 10 or 5°C, either from eclosion or from 8 days after eclosion, the survival rate, on testing, rose with time since transfer and rose faster and higher with a photoperiod of LD 10:14h than with LD16:8h. Flies transferred to 15°C only showed improved ability to survive the test if they were kept in LD 10:14h. When cultured at 18°C to the age of 8 days after eclosion, diapause was terminated in about 30% of females even at LD 10:14h. In these post-diapause females the ability to develop cold-hardiness at lower temperatures was somewhat less than in the diapausing females, but apparently greater than in the non-diapause females. These results suggest that the physiological mechanism which promotes cold-hardiness under a diapause-inducing photoperiod is not directly linked to the process causing reproductive diapause.
In Sapporo, flies from a natural population became tolerant to cold in October when they entered diapause and daily mean temperature fell below 15°C and the light/dark cycle fell below LD 12:12h.     

Effects of photoperiod and temperature on the development and diapause of the bark beetle Ips typographus

Abstract:  Diapause was induced in a Central European population of Ips typographus grown at 20°C when the day length decreased below 16 h [50% diapause incidence occurred in the 14.7:9.3 h L:D (light:dark) regime]. The non-diapausing adults fed on days 2–6 and 10–14 after the ecdysis and swarmed after the second feeding bout with chorionated eggs in the ovaries and sperm in the spermiducts. Neither gonads nor the flight muscles matured and no swarming occurred in the diapausing adults. The development from egg to adult took about 34 days in both 18:6 h (no diapause) and 12:12 h L:D (diapause) regimes, but it was extended by up to 30% without diapause induction when only larvae or pupae were exposed to L:D 12:12 h. Diapause was induced in insects reared at L:D 12:12 h through the last larval and the pupal instars and/or in the adult stage. Temperature ≥ 23°C prevented diapause induction at L:D 12:12 h but diapause occurred at L:D 14:10 h associated with 26:6°C thermoperiod. The effect of thermoperiods on the developmental rate requires further research. Exposure of the non-diapausing adults to 5°C for several days blocked feeding and evoked a diapause-like state, whereas diapausing adults fed and their gonads slowly developed at this temperature. Diapausing adults exposed in forest to low night temperatures and transferred in October to 20°C readily reproduced at 18:6, but not 12:12 h L:D photoperiods. After 2-months at 5°C and darkness, they became insensitive to the photoperiod, matured and most of them also swarmed at 20°C in the 12:12 h L:D regime. In a Scandinavian population, diapause occurred at 18:6 h L:D and was terminated either by exposure to 5°C or by very long photoperiod (L:D 20:4 h) combined with high temperature (23°C).