Monarch butterflies reared under autumn-like conditions have more efficient flight and lower post-flight metabolism

1. Many migratory animals undergo physiological and behavioural changes to prepare for and sustain long-distance movements. Because insect migrations are common and diverse, studies that examine how migratory insects meet the energetic demands of long-distance movements are badly needed. 2. Monarch butterflies (Danaus plexippus) migrate up to 4000 km annually from eastern North America to wintering sites in central Mexico. Autumn generation monarchs undergo physiological and behavioural changes in response to environmental cues to initiate migration. In particular, exposure to cooler temperatures and shorter day lengths in early autumn causes monarchs to enter the hormonally induced state of reproductive diapause. 3. This study examined differences in flight-associated metabolic rate (MR) and flight performance metrics for monarchs experimentally reared under autumn-like conditions (typically experienced before the southward migration) relative to monarchs reared under summer-like conditions. 4. Adult monarchs reared under autumn-like conditions showed lower post-flight MRs, greater flight efficiency, and lower measures of reproductive activity relative to monarchs reared under summer-like conditions. Increases in post-flight metabolism were associated with monarch body weight, age, and flight velocity. 5. These findings suggest that a trans-generational shift in flight energetics is an important component of the monarch's complex migratory syndrome, and that physiological changes that accompany reproductive diapause facilitate energy conservation during flight.     

寄主植物与温度对大豆食心虫滞育期间糖类和脂质含量的影响

【目的】本文探究了大豆食心虫Leguminirora glycinioorella(Mats.)Obraztsov进入滞育以后体内糖类、脂质和水分含量的变化,以及不同寄主植物和滞育诱导期温度对大豆食心虫体内糖类、脂质和水分含量的影响。【方法】通过蒽酮硫酸法和氯仿甲醇分离法测定了滞育前后、以大豆Glycine max(L.)Merr和野大豆Glycine soja Sieb.et Zucc.两种植物为寄主以及不同滞育诱导温度下的大豆食心虫体内脂质、水分、总糖、糖原和海藻糖的含量,结果利用SPSS17.0数据分析软件进行分析。【结果】进入滞育阶段的大豆食心虫体内脂质、总糖、海藻糖和糖原含量显著增加,含水量显著下降。以大豆为寄主的大豆食心虫在滞育阶段体内水分含量、总糖和糖原含量较野大豆寄主的高,海藻糖含量较野大豆的低,差异均显著。不同滞育诱导温度下的大豆食心虫体内生化物质含量基本一致。【结论】滞育前后大豆食心虫体内各种生化物质变化明显,以栽培大豆为寄主和以野大豆为寄主的大豆食心虫体内生化物质含量不同,而滞育诱导温度对大豆食心虫体内生化物质含量影响不大。     

Expression of alternative developmental pathways in the cabbage butterfly,Pieris melete and their differences in life history traits

The seasonal life cycle of the cabbage butterfly, Pieris melete is complicated because there are three options for pupal development: summer diapause, winter diapause, and nondiapause. In the present study, we tested the influence of temperature, day length, and seasonality on the expression of alternative developmental pathways and compared the differences in life history traits between diapausing and directly developing individuals under laboratory and field conditions. The expression of developmental pathway strongly depended on temperature, day length, and seasonality. Low temperatures induced almost all individuals to enter diapause regardless of day length; relatively high temperatures combined with intermediate and longer day lengths resulted in most individuals developing without diapause in the laboratory. The field data revealed that the degree of phenotypic plasticity in relation to developmental pathway was much higher in autumn than in spring. Directly developing individuals showed shorter development times and higher growth rates than did diapausing individuals. The pupal and adult weights for both diapausing and directly developing individuals gradually decreased as rearing temperature increased, with the diapausing individuals being slightly heavier than the directly developing individuals at each temperature. Female body weight was slightly lower than male body weight. The proportional weight losses from pupa to adult were almost the same in diapausing individuals and in directly developing individuals, suggesting that diapause did not affect weight loss at metamorphosis. Our results highlight the importance of the expression of alternative developmental pathways, which not only synchronizes this butterfly's development and reproduction with the growth seasons of the host plants but also exhibits the bet‐hedging tactic against unpredictable risks due to a dynamic environment.     

Host plant mediation of diapause induction in the cabbage beetle, Colaphellus bowringi Baly （Coleoptera： Chrysomelidae）

The cabbage beetle, Colaphellus bowringiBaly （Coleoptera： Chrysomelidae）, is a serious pest of crucifers in China, undergoing an imaginal summer and winter diapause in the soil. The effects of host plants on diapause incidence were tested in the beetle. The ratio of adults entering diapause was significantly low when they fed on the mature leaves of Chinese cabbage Shanghaiqin （Brassica chinesis var communis） than those feeding on Chinese cabbage Suzhouqin （Brassica chinesis var communis）, radish （Raphanus sativus var longipinnatus） and stem mustard （Brassicajuncea vat tumida） at 25℃ combined with 13：11 （L： D） h. Fewer adults entered diapause on young leaves compared to physiologically aged and mature radish leaves at 25℃ combined with 13：11 （L： D） h. The effect of host plant species on diapause induction was also evident under continuously dark rearing conditions or at different photoperiods. These experimental results demonstrate that host plant mediation of diapause induction exists in the cabbage beetle. However, at temperatures ≤20℃ or photoperiods of 16：8 （L： D） h combined with 25℃, all individuals entered diapause regardless of the host plants, indicating that the effects of host plants on diapause induction could be expressed only within a limited range of temperatures and photoperiods.     

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.     

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.     

Photoperiodic,thermal and trophic responses of a predatory ladybird Cheilomenes propinqua

Large geographic range, wide habitat specificity and broad range of prey (including a number of pests of protected crops) suggest that a predatory ladybird Cheilomenes propinqua can be considered as potential agent for biological control in greenhouses. We investigated the influence of day length (10, 12 and 14 hr), temperature (20 and 24°C) and diet (the green peach aphid Myzus persicae and eggs of the grain moth Sitotroga cerealella) on the rate of maturation, fecundity and induction of reproductive diapause in C. propinqua females of a laboratory population originated from Alexandria, Egypt. The proportion of diapausing females (i.e. those with poorly developed ovaries and well-developed fat body) varied from 5% to 70% being higher at short day, low temperature and feeding on the grain moth eggs. This diapause, however, was not very stable: more than 20% of females kept under diapause-inducing conditions started to lay eggs during 110 days, although their pre-oviposition period was about 5 times longer than that of females which matured at the same temperature but at the long day and on the natural diet. Although not very stable, reproductive diapause significantly increased survival of starving females. Such a short-term reproductive diapause can be considered as an adaptation to mild and short-term subtropical winter. The results of our study suggest that C. propinqua mass rearing will be more intensive at the combination of high temperature, natural food (aphids) and long day (14 hr), whereas individuals intended for long-term storage should be reared under moderate temperature (20°C) and short-day (10 hr) conditions and should be fed on factitious food (the grain moth eggs).     

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.     

Sensitivity to juvenile hormone is not reduced in clustering monarch butterflies, Danaus plexippus, in Australia

ABSTRACT. Female monarch butterflies, Danaus plexippus (L.) were collected from clusters in the Sydney region at weekly intervals after cluster formation. They were neck-ligated and injected with juvenile hormone (JH) in order to test for any period of lowered JH sensitivity during the non-reproductive clustering phase. Response was assessed by production of mature oocytes. In all weekly samples, the mean response was equal to or greater than that of newly emerged females reared and maintained in optimal conditions, and was substantially higher than that reported for diapausing monarchs in North American clusters. The result is further evidence for the absence of a true reproductive diapause in New South Wales populations.     

Effects of temperature of adults and eggs on the induction of embryonic diapause in the band-legged ground cricket, Dianemobius nigrofasciatus

Abstract.  Eggs laid by adult female Dianemobius nigrofasciatus , reared under long-day (LD 16 : 8 h, 25 °C) or short-day (LD 12 : 12 h, 25 °C) conditions from the nymphal stage, are kept at several constant temperatures. At 22.5–30.0 °C, eggs laid by long-day adults show lower incidences of diapause than those laid by short-day adults. In both eggs laid by adults under long-day conditions and those under short-day conditions, the higher the temperature at which the eggs are kept, the lower the incidence of diapause. When eggs of long-day adults are exposed to a low-temperature pulse (10 °C, 24 h) on the day of deposition (day 0), the incidence of diapause increases. The low-temperature pulse on day 1 does not increase the incidence of diapause. By contrast, when the eggs of short-day adults are exposed to a high-temperature pulse (35 °C, 24 h) on day 0 or day 1, the incidence of diapause decreases. The temperature pulses on day 0 are more effective at diapause prevention. Staining of diapause eggs by the Feulgen–Rossenbeck method shows that the eggs enter diapause at the blastoderm stage, which is on day 1 or day 2 at 25 °C. The exposure of adults to long days and higher temperatures prevents the eggs from entering diapause. In D. nigrofasciatus , embryonic diapause is controlled by maternal effects, adult photoperiod and temperature, and egg temperature before or at diapause.     

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

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

Feeding insects with a phytopathogenic fungus influences their diapause and population dynamics

Abstract.  1. A facultative mutualistic relationship between a herbivorous moth, Lobesia botrana , and a phytopathogenic fungus, Botrytis cinerea , both damaging the same plant, the vine, has been demonstrated recently. Laboratory and field studies were carried out to determine the influence of the presence of the fungus in food on larvae during the pre- and post-diapause periods and on the global fitness of the insect.
2. During the pre-diapause period, larvae fed on fungus exhibited a higher survival rate and a faster larval development than insects reared without fungus.
3. After the wintering diapause, insects reared with fungus displayed a better synchronisation of adult emergence and an increased fecundity compared with control insects.
4. These results indicate that food containing fungus provides an improved diet, permitting the herbivorous insect to obtain nutrients required to optimise their life-history traits.
5. In addition, the results emphasise the importance of the nutritional quality of larval diet, improved by addition of the fungus, on diapause termination and on the demography of the insect before and after the wintering diapause.     

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

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Time of the season: the effect of host photoperiodism on diapause induction in an insect herbivore,Leptinotarsa decemlineata

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Mating prior to overwintering in the imported willow leaf beetle, Plagiodera versicolora (Coleoptera: Chrysomelidae)

Abstract. 1. We report on the mating status of overwintering adults of the imported leaf beetle, Plagiodera versicolora Laicharting. This species overwinters as adults, under the bark of willow trees.
2. Individuals were collected both in the autumn, just after initiation of diapause, and in the spring, just before natural termination of diapause. From the autumn sample, we discovered that diapause can be terminated by husbanding the adults at 4°C for 4 weeks.
3. Most females lay fertile eggs upon the termination of diapause; however, a significant number of females lay non-fertile eggs after overwintering in the field.
4. Many females mate with more than one male before entering diapause. Progeny analysis using allozyme genetic markers shows that, on average, females who are fertile upon termination of diapause utilize sperm from 1.2 males to fertilize their eggs.     

Life-History Variation inChrysoperla carnea:Implications for Rearing and Storing a Mexican Population

A population ofChrysoperla carneafrom San Pedro, Coahuila, Mexico, showed phenological responses that differ from those of previously studied North American biotypes. Short daylengths that induce diapause in other biotypes resulted in low incidences of diapause in the San Pedro population. Diapause induction required a decrease in daylength (from long to short days). Constant long days produced variable reproductive responses, whereas an increase in daylength (from short to long day) resulted in high incidences of reproduction. Providing prey to adults that had been reared under short days consistently increased the incidences of reproduction. Storage of either diapausing or nondiapausing adults at low temperature (5°C) for 13 weeks yielded better survivorship and reproductive performance than in unstored (24°C) controls. For efficient production, the San Pedro population ofC. carneamay be reared either with an increase in daylength or with the regular intervention of a storage period. To induce diapause in individuals destined for storage, larvae may be reared under long-day photoperiod and the prepupae transferred to short days.     

Phenology of weight, moisture and energy reserves of Australian monarch butterflies, Danaus plexippus

Abstract. 1. In the Sydney area of New South Wales, dry weights of reproductive monarch butterflies averaged 156 mg and were higher during winter than in other seasons. Dry weights of non-reproductive monarchs ranged from 216 to 324 mg and declined by 15–25% during over wintering.
2. Fat in reproductive butterflies ranged from 0.009 to 0.017g/0.1g dry weight and was lower during winter than in other seasons. Fat content of non-reproductive monarchs was higher (0.019–0.037 g/0.1g) and declined by 24–51% during over wintering.
3. Lean dry weights of reproductive monarchs were lower than those of non-reproductive individuals. Lean dry weights of non-reproductive butterflies increased rapidly at the beginning of over wintering and remained high throughout the winter. Analysis of protein content indicated the higher lean weight of non-reproductive monarchs was due to greater protein levels.
4. Moisture content of monarchs did not vary with season or reproductive status and appeared to be correlated with ambient humidity.
5. Non-reproductive monarch butterflies in New South Wales adjust bio chemically during over wintering. Energy reserve dynamics of these butterflies are comparable to those that occur in non-reproductive monarchs in North America.     

Diapause affects cuticular hydrocarbon composition and mating behavior of both sexes in Drosophila montana

Envipnmental cues,mainly photoperiod and temperature,are known to control female adult reproductive diapause in several insect species.Diapause enhances female survival during adverse conditions and postpones progeny production to the favorable season.Male diapause(a reversible inability to inseminate receptive females)has been studied much less than female diapause.However,if the males maximized their chances to fertilize females while minimizing their energy expenditure,they would be expected to be in diapause at the same time as females.We investigated Drosophila montana male mating bchavior under short-day conditions that induce diapause in females and found the males to be reproductively inactive.We also found that males reared under long-day conditions(reproducing individuals)court reproducing postdiapause fermales,but not diapausing ones.The diapausing fies of both sexes had more long-chain and less short-chain hydrocarbons on their cuticle than the reproducing ones,which presumably increase their survival under stressful conditions,but at the same time decrease their attractiveness.Our study shows that the mating behavior of females and males is well coordinated during and afier overwintering and it also gives support to the dual role of insect cuticular hydrocarbons in adaptation and mate choice.     

Host plant and latitude-related diapause variation in Rhagoletis pomonella: a test for multifaceted life history adaptation on different stages of diapause development

Variation in the overwintering pupal diapause of Rhagoletis pomonella appears to adapt sympatric populations of the fly to seasonal differences in the fruiting times of their host plants, generating ecological reproductive isolation. Here, we investigate what aspects of diapause development are differentially affected (1) by comparing the propensities of apple vs. hawthorn-infesting host races of R. pomonella to forgo an initially deep diapause and directly develop into adults, and (2) by determining the chronological order that R. pomonella races and sibling species break diapause and eclose when reared under standardized environmental conditions. The results imply that factors affecting initial diapause depth (and/or differential mortality during the prewintering period) and those determining the timing of diapause termination or rates of post-diapause development are both under differential selection and are to some degree genetically uncoupled in flies. The modular nature of diapause life history adaptation in Rhagoletis suggests that phenology may involve multiple genetic changes and represent a stronger ecological barrier separating phytophagous specialists than is generally appreciated.     

A true summer diapause induced by high temperatures in the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae)

Summer diapause in the cotton bollworm, Helicoverpa armigera (Hübner), which prolongs the pupal stage, particularly in males, is induced by high temperatures. In the laboratory, summer-diapausing pupae of H. armigera were induced at high temperatures (33-39 degrees C) with a photoperiod of LD8:16; winter-diapausing and non-diapausing pupae, cultured at 20 degrees C with a photoperiod of LD8:16 and at 27 degrees C, LD16:8, respectively, acted as a control. Retention time of eye spots, weight, and lipid and glycogen levels were compared. At high temperatures, both body weight and energy storage capacity were much higher in summer-diapausing pupae than in non-diapausing pupae reared at 33-39 degrees C. At temperatures (>33 degrees C) high enough to maintain summer diapause, the eye spots of summer-diapausing pupae did not move during the 30-day experiment. However, eye spots of summer-diapausing pupae placed at 30 degrees C began to move about 10 days after they were transferred, significantly later than in non-diapausing pupae reared at 33-39 degrees C or non-diapausing pupae reared at 27 degrees C, which initiated eye spot movement 2 days after pupation. The differences in retention time of eye spots between summer- and winter-diapausing pupae shows that winter diapause is more intense than summer diapause in this insect. The weight loss, and lipid and glycogen metabolism curves indicate that the summer-diapausing pupae's metabolism is very low. We conclude that summer diapause in the cotton bollworm is a true diapause and that the summer diapause enables the cotton bollworm to withstand the high temperatures of summer.