Differences in photoperiod‐induced diapause plasticity among different populations of the bark beetle Ips typographus and its predator Thanasimus formicarius

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Reproductive diapause and the bacterial symbiosis in the sycamore aphid Drepanosiphum platanoidis

1. Parthenogenetic adults of the sycamore aphid Drepanosiphum platanoidis exhibited reproductive diapause for 4–6 weeks in the summer. 2. The diapausing aphids had small gonads (accounting for just 13% of the total aphid protein content) bearing small (< 0.5 mm long) and developmentally immature embryos. There was no evidence of embryo resorption. 3. The diapausing aphids had significantly depressed essential amino acid content and elevated glutamine content, relative to reproductively active D. platanoidis. 4. The reproductive characteristics and amino acid titres of the diapausing aphids resemble those of aphids lacking functional symbiotic bacteria. Uncoupling of maternal and embryo growth and suppression of bacterial function are proposed as key elements in the diapause of D. platanoidis.     

Seasonal variation in generation time, diapause and cold hardiness in a central Ohio population of the flesh fly, Sarcophaga bullata

Abstract.

• 1 Generation time, diapause phenology and cold tolerance of the flesh fly, Sarcophaga bullata, were examined under confined natural conditions in central Ohio. In this locality, the fly can complete a maximum of four generations annually.
• 2 Very few pupae entered diapause in the first and second generations (May to July in 1988). In the third generation (August) 37% of the pupae entered an overwintering diapause, as did all pupae from the fourth generation (September).
• 3 The adult eclosion date in the spring and annual generation time can be predicted accurately from degree day data.
• 4 Cold tolerance of the field-overwintering portion of the population was high. After 30 days under field conditions, diapausing pupae readily survived a 7-day exposure to — 17°C. Glycerol appears to be the major cryoprotectant in S.bullata, and glycerol concentrations in the field population (95–142 mm ) remained high throughout the winter.
• 5 In contrast, diapausing flies reared under laboratory conditions (20°C, 12:12 LD) were less cold tolerant, and glycerol concentrations were lower (6.9–21.2 mm ). Field conditions thus promote the acquisition of high levels of cold tolerance, presumably as a consequence of the accumulation of higher concentrations of glycerol.
• 6 In spite of differences in the cold tolerance of laboratory and field flies, the supercooling points of the two groups of flies were nearly the same, thus implying that the supercooling point is not a good indicator of cold tolerance.

     

Induction of reproductive diapause in Habrobracon hebetor (Hymenoptera: Braconidae) when reared at different photoperiods at low temperatures

Development of the parasitoid Habrobracon hebetor (Say) (Hymenoptera: Braconidae) at low temperatures was determined to identify rearing conditions that might result in adults that were in reproductive diapause. Diapausing adults would be expected to survive cold storage longer than nondiapausing adults for use in biological control programs. Only a few eggs were found in the ovaries when H. hebetor females were reared during the immature stages at 17.5 and 20°C with a 16-h photoperiod, and the ovaries were poorly developed and contained no eggs when females were reared with a 10-h photoperiod in these low temperatures. Rearing H. hebetor at 17.5 and 20°C did not result in diapause of immature stages, but did appear to result in possible adult reproductive diapause when the immature stages were reared with a 10-h photoperiod. Females reared during the immature stages at 17.5°C with a10-h photoperiod lived longer and took longer to lay their first eggs and to lay 50% of their eggs than those females reared at 17.5°C with a16-h photoperiod. Females reared during the immature stages at 20°C with a10-h photoperiod took longer to lay their first eggs and to lay 50% of their eggs, and they had a lower respiration rate, than those females reared at 20°C with a16-h photoperiod. Females that were reared in conditions that appeared to induce reproductive diapause resumed oviposition and their respiration rate increased soon after being transferred to a higher temperature (27.5°C). Thus, females reared at a 10-h photoperiod at 17.5 and 20°C appear to enter reproductive diapause.     

Role of a parasitoid with a low fecundity,Praestochrysis shanghaiensis (Hymenoptera: Chrysididae), in the population dynamics of its host

1. Analysis of life tables of the oriental moth, Monema flavescens, obtained for 8 generations over 4 years, disclosed that the cocoon parasitoid, Praestochrysis shanghaiensis, acted as a density-disruptive factor.
2. The density of the host cocoon remained stable (max./min.=3.2), whereas that of the host adult varied (max./min.=14.3) although both showed similar fluctation patterns.
3. Stability of the host population was associated with the density-dependence in the ratio of first generation cocoons to overwintered generation moths, which was the key factor for the rate of change throughout the year. Chrysidid parasitism among the first generation cocoons ranged from 37.7 to 70.1%, and that among the second generation cocoons from 16.7 to 63.2%, each showing an inverse density-dependence and acting as the main determinant (key-factor) of the between-year variation in the density of the adult moths.
4. The density-dependence of the rate of change from overwintered generation adults to first generation cocoons was so strong that the parasitism on the second generation hosts had not effect on the cocoon density of the first generation. On the other hand, the density-dependence of the rate of change from first generation adults to second generation cocoons was weak, and the parasitism on the first generation hosts became the key factor for the between-year variation of the second generation cocoons.
5. It is suggested that the stability of the parasitoid-host system will be disrupted without three parasitism-restricting factors: asynchrony in the parasitoid attack on the second generation hosts, high mortality among parasitoid larvae of the second generation, and the high proportion of those first generation parasitoids that enter diapause. These factors are considered to be effective only in cooler parts of the distribution of the parasitoid.

     

Resource tracking in aphids: programmed reproductive strategies anticipate seasonal trends in habitat quality

Most aphids show cyclical parthenogenesis, have short generation times and complete several generations each season. Two hypotheses have been proposed to account for the adaptive seasonal trends observed in the reproductive strategies of aphids. Firstly, individuals of each generation modify their reproductive strategy in direct response to the conditions they experience during their development. Secondly, the reproductive strategies of the different generations are to a large extent programmed and anticipate seasonal trends in habitat quality. These hypotheses were tested by rearing individuals of three generations of the host-alternating willow-carrot aphid, Cavariella aegopodii, on both willow and carrot. This revealed that the way this aphid allocates resources to gonads and lipid reserves is independent of an aphid's weight and the host plant on which it is reared. In addition each generation shows a specific relationship between offspring size and adult size, which tends to keep the absolute investment in individual offspring relatively constant from generation to generation, inspite of the big differences in adult size between generations. That is, through programmed allometric engineering aphids anticipate the predictable seasonal trends in habitat quality and so more closely track their resources, investing relatively more in gonads when food quality is high and relatively more in lipoidal reserves when food quality is poor. Received: 25 August 1997 / Accepted: 19 October 1997     

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.     

Impact of transgenic cotton plants on a non-target pest, Aphis gossypii Glover

Abstract.  1. Transgenic crops have shown great promise for the control of target pest insects, but in some cases they can also influence non-target species. This study investigated the impact of Bt and Bt+CpTI transgenic cottons on the non-target cotton aphid, Aphis gossypii Glover, by comparing life-table parameters, feeding behaviour, and the fluctuating asymmetry of morphological traits of aphids reared on transgenic cotton and those on untransformed control plants.
2. Aphids on the Bt+CpTI cotton showed a shorter reproductive duration and maximum lifespan, lower survival rates and potential maximum fecundity, and an earlier occurrence of peak daily mortality in the first or second generation. However, the aphid population soon developed fitness and overcame the negative effect in the second or third generation. The aphids on the Bt cotton had significantly longer reproductive durations in the first generation, higher survival rates in the third generation, and apparently larger potential maximum fecundity in all three generations.
3. The percentages of accumulated duration of feeding waveforms E1 and E2 were significantly lower in aphids on the Bt+CpTI cotton than in those on the Bt or control cotton, whereas the frequencies of moving and finding feeding sites, and probe behaviour were significantly higher.
4. Fluctuating asymmetry in three morphological characters of aphids reared on transgenic and control cotton was detected. The fluctuating asymmetry value of the third segment of antenna in aphids on Bt+CpTI cotton was significantly higher than that of aphids on Bt or control cotton. Based on the fluctuating asymmetry value, the stress of cotton on the aphids could be ranked as Bt+CpTI cotton > Bt cotton > control cotton.     

Habitat quality and reproductive investment in aphids

ABSTRACT.

• 1 Genetically identical vetch aphids, Megoura viciae (Buckton), showed a weak but significant positive relationship between reproductive investment (number of ovarioles) and weight, here used as an indicator of habitat quality.
• 2 Apterae reared on poor quality plants showed the same range in numbers (twelve to twenty-two) but on average fewer ovarioles than those reared on high quality plants.
• 3 The variability in ovariole number was determined prior to birth.
• 4 Under constant conditions the age of a mother does not affect the range of ovariole numbers or the proportion of her offspring in each ovariole class.
• 5 Apterae reared on poor quality host plants gave birth to proportionately fewer offspring with a high ovariole number than apterae reared on high quality plants.
• 6 Winged individuals gave birth to proportionately more offspring with a high number of ovarioles than apterae.
• 7 The control and ecological significance of the intramorph variation in reproductive tactics shown by many species of aphids is discussed in the light of these results.

     

Diapause induction and termination in a commonly univoltine leaf beetle (Phratora vulgatissima)

Abstract The leaf beetle Phratora vulgatissima (Linnaeus 1758) is commonly univoltine in south‐central Sweden but may sometimes initiate a partial second generation. The current study was set out to investigate under what abiotic conditions the beetles initiate a second generation. Using climate chamber experiments, the beetles were shown to have a facultative reproductive diapause induced by declining day‐length. The critical day‐length (CDL) for diapause induction was estimated to be 18 h and 10 min. In the field, first‐generation beetles developing to adulthood before August in 2009 became reproductively active and produced a second generation, but most individuals emerged later and were in reproductive diapause. P. vulgatissima overwinter as adults and diapause was shown to be maintained until mid‐winter in 2008/2009. The cumulative temperature requirement for oviposition after diapause termination was estimated to be 222 day‐degrees with a 5.5°C temperature threshold. Three different day‐degree models that were developed to predict the phenology of female oviposition in the spring were validated by comparing model results with field data on the timing of oviposition in previous years. The study suggests that P. vulgatissima may initiate a second generation in Sweden if development of the first generation is completed before August. Warmer spring and summer temperatures due to ongoing climate change may cause advanced insect phenology and faster completion of insect life‐cycles at northern latitudes, which will affect the proportion of insects that initiate a second generation.     

Dispersion of flightless adults of the Asian lady beetle, Harmonia axyridis, in greenhouses containing cucumbers infested with the aphid Aphis gossypii: effect of the presence of conspecific larvae

Nezara viridula (L.) (Heteroptera: Pentatomidae) recently expanded its distribution range in Japan and reached Osaka. In the southern temperate zone, the species overwinters in the adult stage and reproductive diapause is associated with a body colour change from green to russet. In Osaka, the reproductive diapause is only induced in September–October and nymphs from late egg masses are destined to die during winter. However, the fate of adults emerging late in the season remained unknown. Survival, body colour change, and post‐diapause reproduction were studied under quasi‐natural conditions in Osaka in 1999–2000 in those adults that attained adulthood as late as in November. Two experimental cohorts were used: in the Outdoor cohort, insects were reared outdoors starting in their second instar (28 September); in the Laboratory cohort, nymphs and subsequently adults were reared from the same day under diapause‐inducing conditions (L10:D14 at 25 °C), then acclimatized (5 days at 20 °C and 5 days at 15 °C; L10:D14) and transferred outdoors on 1 December. Adults in both cohorts did not reproduce in autumn and survived the winter with a low mortality. More than 20% of adults in the Outdoor cohort failed to change body colour from green to russet during winter, apparently because of the low ambient temperature, suggesting that the environmental conditions required for colour change do not completely coincide with those required for diapause induction, and that the colour of the adults is not always a reliable indicator of diapause in this species. After overwintering, females from the Outdoor cohort produced significantly fewer egg masses and eggs and had a significantly shorter period of oviposition than females that entered diapause under short‐day conditions in the Laboratory cohort. Thus, if progeny from the late egg masses attain adulthood late in the autumn, these adults have high chances of successful overwintering, but their reproductive output after the winter diapause is significantly reduced.     

Host‐plant quality adaptively affects the diapause threshold: evidence from leaf beetles in willow plantations

1. Voltinism of herbivorous insects can vary depending on environmental conditions. The leaf beetle Phratora vulgatissima L. is univoltine in Sweden but will sometimes initiate a second generation in short‐rotation coppice (SRC) willow plantations. 2. The study investigated whether increased voltinism by P. vulgatissima in plantations can be explained by (i) rapid life‐cycle development allowing two generations, or (ii) postponed diapause induction on coppiced willows. 3. In the field, no difference was found in the phenology or development of first‐generation broods between plantations (S. viminalis) and natural willow habitats (S. cinerea). However, the induction of diapause occurred 1–2 weeks later in SRC willow plantations. 4. Laboratory experiments indicated no genetic difference in the critical day‐length for diapause induction between beetles originating from plantations and natural habitats. Development time was unaffected by host‐plant quality but critical day‐length was prolonged by almost an hour when the beetles were reared on a non‐preferred willow species (S. phylicifolia). When reared on new leaves from re‐sprouting shoots of recently coppiced willow plants, diapause incidence was significantly less than when the beetles were reared on mature leaves from uncoppiced plants. 5. The study suggests that P. vulgatissima has a plastic diapause threshold influenced by host‐plant quality. The use of host‐plant quality as a diapause‐inducing stimulus is likely to be adaptive in cases where food resources are unpredictable, such as when new host‐plant tissue is produced after a disturbance. SRC willows may allow two beetle generations due to longer growing seasons of coppiced plants that grow vigorously.     

Heterodynamic processes in Coccinella septempunctata L. (Coccinellidae: Coleoptera): a mini review

Among ladybirds, diapause has evidently been most studied in Coccinella septempunctata L. adults. Coccinella septempunctata (C‐7), the seven spot, has attracted attention both in Europe and the USA for being heterogeneous as to the induction of diapause. A polyphenic character of diapause is a prominent feature in C. septempunctata and this phenomenon generally and often generates voltinism heterogeneity within populations. A greater part of the C‐7 population shows an obligatory univoltine cycle, whereas a smaller proportion is facultatively polyvoltine. Coccinella s. bruckii in Japan is bivoltine, in which the first generation of adults aestivate while the second generation hibernates. This paper reviews this heterodynamic cycle and its importance in the life history of C‐7, as reported from different regions of the world. Heterogeneous voltinism and diapause smooth the progress of a portion of C‐7 populations to take benefit of aphids well into late summer and autumn. Understanding such variation and plasticity in the life history of this species can help in organizing proper biological control attempts using it as a biocontrol agent.     

First evidence for photoperiodic regulation of the life cycle in a millipede species, Polydesmus angustus (Diplopoda: Polydesmidae)

The hypothesis that the periods of dormancy previously described in the millipede Polydesmus angustus may be photoperiodically induced diapauses was tested experimentally. In this species, biennial individuals exhibit two successive periods of dormancy: aestivation in the penultimate stadium (stadium VII) and reproductive dormancy in the adults, which emerge in autumn. It was first established that the reproductive dormancy is not a thermally controlled state of quiescence. When adults emerging in autumn were kept at 16 °C under natural photoperiod, their reproduction was delayed for several months in comparison with adults emerging in spring at similar temperatures. This indicates that the reproductive dormancy begins with a period of diapause. Further experiments provided evidence of a photoperiodic induction of the adult diapause. When millipedes were reared under short day length (L:D 12:12 h) throughout their development, they required more time to reproduce than millipedes reared under long day length (L:D 16:8 h) at the same temperatures. Photoperiod influenced reproduction in females, but no significant effects were detected in adult males. On the other hand, stadium VII was markedly longer at L:D 16:8 h than at L:D 12:12 h in both sexes, which strongly suggests that aestivation is also induced by photoperiod. However, the effects on the duration of stadium VII varied among individuals, some of which showed no response to long days. This study is the first to document photoperiodic regulation of the life cycle in the class Diplopoda, a trait common in other classes of terrestrial arthropods.     

Reproductive biology of the arctic collembolan Hypogastrura tullbergi

Terrestrial organisms of the Arctic are faced with strong climatic fluctuations. Predictable seasonality with cold/long winters and short/cool summers are combined with unpredictable between and within year variation. This indicates that various selection pressures act on the reproductive strategies of the populations. The arctic collembolan Hypogastrura tullbergi reproduces in a short period following snow melt. Hatching occurs in late summer, the animals grow to adult size within their second summer and reproduce for the first time in the beginning of their third summer. We performed several experiments to determine the reproductive investment and proximate mechanism that regulate timing and duration of reproduction. We found that H. tullbergi entered a reproductive diapause when reared at constant temperature, a diapause that was terminated by a cold exposure (winter). Surprisingly, cold exposure of small juveniles may also prevent development of a reproductive diapause in adults. Thus, the life-cycle normally spanning 2 yr can potentially be reduced to one year if the overwintering juveniles reach maturity before the end of the reproductive period in the field. After termination of the diapause, the animals reproduced up to 3 times during a period of 6 weeks at 15°C. This reproductive period was considerably longer (measured in degree days) than the one observed in the field. Our results suggested that temperature quiescence, i.e. the inability to reproduce under a certain temperature threshold, may adjust the termination of the reproductive period with current temperature before a new diapause occurs in late summer. The cost of reproduction was low and suggests that it can be adaptive to spread reproduction over more than one year. The results are discussed in relation to the arctic climate and strategies favoured by unpredictable and predictable (seasonal) variations in the environment. The present study forms part of a larger investigation on population dynamics and life history strategies of H. tullbergi from the Arctic.     

Dynamics of the Photo-Trophic Termination of Reproductive Diapause in Females of the Multicolored Asian Ladybird <Emphasis Type="Italic">Harmonia axyridis</Emphasis> (Pallas) (Coleoptera,Coccinellidae)

Females of certain aphidophagous ladybirds in the absence of natural protein food (aphids) enter reproductive diapause. Reactivation of diapausing beetles is possible only after consuming the food which is suitable for oogenesis of females and development of larvae. The influence of diet and photoperiod on the dynamics of weight and on the rate of reproductive maturation of reactivating females of Harmonia axyridis was studied under laboratory conditions. The experiments were conducted at combinations of two day lengths (12 and 18 h) with 6 trophic regimes with the following mean numbers of daily consumed aphids: 0 (aphids were absent over the whole experiment), 0.1 (1 aphid was provided every 10 days), 0.5 (1 aphid every second day), 1 (1 aphid every day), 10 (10 aphids every day), and 50 (about 50 aphids every day). Judging from the state of ovarian development estimated by dissection 20 days after the beginning of the experiment, the threshold of the trophic termination of reproductive diapause under the long day conditions lies between the regimes of 0.1 and 0.5 aphids per day, which is approximately equal to the earlier investigated threshold of the trophic induction of diapause. Short day slowed reactivation and has increased the threshold of the trophic response up to 1-10 aphids per day. Based on these data, we conclude that under natural conditions females which entered reproductive diapause because of the absence of aphids are capable of reactivation in the presence of even a minimal amount of natural protein food. Moreover, in spring and summer (when the probability of the increase in aphid abundance is relatively high) reactivation starts at lower prey population density than in autumn, when the appearance of the stable aphid colony (which is necessary to complete the development of the emerged larvae) is less probable.     

Environmental Cues,Endocrine Factors,and Reproductive Diapause in Male Insects

Environmental cues, mostly photoperiod and temperature, mediated by effects on the neuroendocrine system, control reproductive diapause in female insects. Arrest of oocyte development characterizes female reproductive diapause, which has two major adaptive functions: It improves chances of survival during unfavorable season(s), and/or it confines oviposition to that period of the year that is optimal for survival of the eggs and progeny. Although reproductive diapause is less well studied in male insects, there may be no sex-dependent differences in regard to the first of these functions. The second one, however, is not valid for the male; instead, selection pressure directs the male's reproductive strategy toward maximum chances of fertilization of the female's eggs with minimum waste of energy. Therefore, in species with female reproductive diapause, the males may or may not exhibit diapause, but if they do, their diapause must be adapted to that existing in conspecific females. Male reproductive diapause is defined as a reversible state of inability of the male to inseminate receptive females. In relation to reproductive diapause, there are several patterns of coadaptations between male reproductive strategy and timing of female receptivity, (a) In some insects, the females are receptive in the early part of their diapause; mating occurs during this period and there is no diapause in the male. The male dies shortly after copulation and the female stores the sperms to fertilize the eggs that develop after termination of the female's diapause, (b) In some species, as in the grasshopper Anacridium aegyptium, females are receptive during diapause; though oocyte development is arrested, copulation occurs and the stored sperms fertilize the eggs when the female's diapause ends. Males were claimed to have no diapause, but recent studies have revealed the presence of a reproductive diapause in a proportion of the males. This and other cases show that female receptivity during reproductive diapause may or may not be accompanied by male reproductive diapause. If there is a reproductive diapause in the male, it is controlled by the same endocrine mechanism, the corpora allata (CA), as in the females, (c) In many species females are refractory during their diapause. In these cases, males exhibit reproductive diapause, which may be light, as in the beetle Oulema melanopus, or well established, as in certain grasshoppers, butterflies, and beetles. In the latter cases, male diapause is controlled by similar environmental cues (photoperiod, temperature) and by the same intrinsic mechanism (neuroendocrine system, especially CA) as female diapause. Nevertheless, male diapause is less intense; the environmental cues leading to its termination are less complex and/or less extreme, so male diapause terminates before that of the females. Presumably, male diapause is under two antagonistic selection pressures: A male should not waste energy by courting dia-pausing refractory females, but he should be ready to copulate as soon as the females become receptive, otherwise he may lose in the competition between males for females. Some further strategies, which do not seem to fit the above patterns, are also outlined.     

Differentiation in life cycle of sympatric populations of two forms of Hyphantria moth in central Missouri

Wing patterns of Hyphantria adult male moths collected in central Missouri were examined throughout the breeding season. Three major peaks of adult flight were observed: the first peak consisted mainly of adults with spotted wings, while the second and third peaks consisted of immaculate adults. Black‐headed larvae appeared in the field following the first major peak of moth flight, and red‐headed larvae appeared in the field following the second peak. Sympatric red‐headed and black‐headed forms were collected in the field and subsequently reared on an artificial diet under conditions of 16 h light : 8 h dark (LD 16:8) at 25°C. The larval period of the black‐headed form was shorter than the red‐headed, whereas the pupal period of the black‐headed form was longer than the red‐headed. Pupal development is retarded in some individuals at high temperatures in the black‐headed form. Photoperiodic response curves for pupal diapause were different between the two forms. The critical photoperiod for pupal diapause was 15 h 10 min in the red‐headed form, which was longer than that for the black‐headed form (14 h 40 min). The two forms responded to shifts in photoperiod differently. These developmental responses temporally separate the two forms in the field; the red‐headed and black‐headed forms represent a set of adaptations favoring univoltinism and bivoltinism, respectively. Red‐headed larvae fed mainly at night, while the black‐headed larvae fed without a clear day–night rhythm. Nocturnal feeding in the red‐headed form is adaptive to protection against predation, but fails to fully utilize heat units and thus to produce a second generation.     

双季稻区稻水象甲一代成虫的滞育

对浙江省双季稻区稻水象甲（ＬｉｓｏｒｈｏｐｔｒｕｓｏｒｙｚｏｐｈｉｌｕｓＫｕｓｃｈｅｌ）夏季种群的研究表明,光照和温度可排除在一代成虫夏季滞育的诱导因子之外,而不同的食料条件是引发滞育的主导因素．在自然状态下,即使是强迫滞留,早稻田内的一代成虫也不转入正常发育,但取食稻秧能使新羽一代成虫和稻田内外处于生殖滞育状态的一代成虫的卵巢正常发育,并能解除已进入夏蛰的一代成虫的生殖滞育;同时,不论是新羽成虫还是夏蛰成虫,取食稻秧后卵巢和飞行肌同时发育;若这种食料条件能持续下去,则飞行肌消解而卵巢发育产卵．     

Developmental and phenotypic responses to photoperiod in uni-and bivoltine Pieris napi in California

• 1 Coastal Californian Pieris napi are facultatively bivoltine, with two seasonal phenotypes, whereas inland populations from the Inner Coast Ranges and Sierra Nevada are univoltine and monophenic.
• 2 When reared under continuous light at 25d̀C both coastal and inland stocks produce about 50% diapause pupae, which give rise to vernal-phenotype adults. Non-diapause pupae of all stocks give rise to summer-phenotype adults, even though this phenotype does not exist in the wild in univoltine populations.
• 3 Univoltinism, which implies developmental suppression of the summer phenotype, is interpreted as a derivative from multivoltinism and an adaptation to host plant phenology.