Control of Diapause by Acidic pH and Ammonium Accumulation in the Hemolymph of Antarctic Copepods

Life-cycles of polar herbivorous copepods are characterised by seasonal/ontogenetic vertical migrations and diapause to survive periods of food shortage during the long winter season. However, the triggers of vertical migration and diapause are still far from being understood. In this study, we test the hypothesis that acidic pH and the accumulation of ammonium (NH₄ ⁺) in the hemolymph contribute to the control of diapause in certain Antarctic copepod species. In a recent study, it was already hypothesized that the replacement of heavy ions by ammonium is necessary for diapausing copepods to achieve neutral buoyancy at overwintering depth. The current article extends the hypothesis of ammonium-aided buoyancy by highlighting recent findings of low pH values in the hemolymph of diapausing copepods with elevated ammonium concentrations. Since ammonia (NH₃) is toxic to most organisms, a low hemolymph pH is required to maintain ammonium in the less toxic ionized form (NH₄ ⁺). Recognizing that low pH values are a relevant factor reducing metabolic rate in other marine invertebrates, the low pH values found in overwintering copepods might not only be a precondition for ammonium accumulation, but in addition, it may insure metabolic depression throughout diapause.     

Climatic conditions, diapause and migration in a troglophile caddisfly

1. Factors explaining the end of diapause include environmental conditions such as daily photoperiod, temperature and humidity. However, because all these factors are almost constant inside deep caves, they would hardly affect diapause termination in many animal taxa that use such habitats in which to aestivate or hibernate (such as bats and some insects, snakes or frogs).
2. An innate biological signal, based on body reserves (mass), could determine the end of diapause in cave-dwelling animals. Another possibility is that the diapausing animals may use as a stimulus convective air circulation, produced by a fall in temperature outside the cave to a value below that inside (i.e. temperature inversions). Here, we explored these two non-exclusive hypotheses explaining the end of diapause and the start of migration in the caddis Mesophylax aspersus . This species aestivates as an adult in caves, as a physiological adaptation to seasonal drought and stream drying.
3. The variation in body mass of individuals entering and leaving the cave was similar, suggesting no role for the 'reserve level' hypothesis in breaking diapause. However, the onset of a temperature inversion in autumn coincided with the breaking of diapause followed after a few days by the migration of the caddisflies.
4. We conclude that the seasonal air currents, produced by thermal inversions, could be the migration stimulus for caddisflies and many other animals that spend a period of diapause in caves.     

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.     

渤海中华哲水蚤(Calanus sinicus)和小拟哲水蚤(Paracalanus Parvus)种群垂直移动生态适应性模型研究

用最优化适合度种群生态模型模拟研究了中华哲水蚤和小拟哲水蚤种群在渤海生态系统中的垂直移动模式。生命周期、世代时间、繁殖力和产卵量是决定这两种桡足类垂直移动的主要因素。在中华哲水蚤生活史类型中,无水平流动或水平流动速率很小时,昼夜的垂直移动能使其获得最优的生态适合度。水流的增加或方向的改变会使其在底层的停留时间增加。在小拟哲水蚤生活史类型中,随着水流的增加及其随之而来的损失率增高,其垂直移动的倾向逐步提高。     

昆虫延长滞育的研究

持续时间达1年以上的滞育,称为延长滞育,延长滞育的诱导、维持、解除均不同于简单滞育(滞育期短于1年)。本论文系统阐述了昆虫延长滞育的类型、滞育诱导及解除的环境因子、延长滞育物候学和生物学特性,延长滞育的遗传学及延长滞育的生态学意义。延长滞育是昆虫生活史的重要组成部分,是一种普遍现象。     

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.     

Effects of photoperiod and temperature on diapause induction in Conogethes punctiferalis (Lepidoptera: Pyralidae)

The yellow peach moth, Conogethes punctiferalis (Guenée), a multivoltine species that overwinters as diapausing larvae, is one of the most serious insect pests on maize in China. Effect of photoperiod and temperature on larval diapause was examined under empirical laboratory conditions. Short‐day treatments caused larval diapause at 25°C, and the critical photoperiod was between 12 and 13 h (or 12 h 51 min) light per day. No sensitive instar was identified for diapause induction under alternated short‐ (L : D 11 : 13 h) and long‐day (L : D 14 : 10 h) treatments at different larval stages. However, accumulative treatment of three instars and 10 d under short‐day treatment was required for the induction of 50% larval diapause. All larvae entered diapause at 20°C, whereas less than 3% did so at 30°C, irrespective of the long‐ or short‐day treatment. Furthermore, under the short‐day treatment, more than 90% of larvae went into diapause with temperatures ≤ 25°C, but less than 17% did so at 28°C. In contrast, under the long‐day treatment, less than 19% of larvae went into diapause with temperatures ≥ 23°C. The forward shift (5°C) of critical temperature under the long‐day regime demonstrated the compensatory effect of temperature and photoperiod on diapause induction. In conclusion, C. punctiferalis had a temperature‐dependent type I photoperiodic diapause response; there was no sensitive instar for diapause determination, but the photoperiodic accumulation time countermeasures both of the short‐day cycles and the number of instars exposed, and the photoperiodic diapause response, was a temperature‐compensated phenomenon.     

Vertical distribution and diel migration of rotifers in a Parana River floodplain lake

The vertical distribution of zooplankton rotifers in the open waters of Laguna El Tigre was investigated. Rotifers showed a relatively uniform distribution throughout the water column. This pattern of distribution was maintained during the year and did not show variations in relation to hydrologic phases of inundation and isolation of the lake. Diel vertical migration of rotifers from the limnetic and the littoral area was investigated too. In littoral area rotifers exhibited a reverse migration, whereas in the limnetic the movements were less conspicuous. Horizontal migration was observed too, and there were interactions between horizontal and vertical distribution. Predation and competition offer a possible explanation.     

Development of genetic differentiation and postzygotic isolation in experimental metapopulations of spider mites

We studied the development of genetic differentiation and postzygotic isolation in experimental metapopulations of the two-spotted spider mite, Tetranychus urticae Koch. A genetically diverse starter population was made by allowing six inbred sublines to interbreed. Then three migration patterns were tested: no migration, or one or three immigrants per subpopulation per generation. Variations in four traits were investigated: allozymes, acaricide resistance, diapause, and hatchability. In the allozymes, acaricide resistance, and diapause, genetic variation among subpopulations became high in metapopulations with no migration, but not in the others, which showed that one immigrant is enough to prevent genetic differentiation. Hatchability, which was decreased by interbreeding among the six sublines, gradually recovered in succeeding generations. In metapopulations with no migration, hatchability was reduced again after in-migration at the 15th generation. Different karyotypes or coadapted gene complexes can survive in different subpopulations by genetic drift, and both Wolbachia-infected and -noninfected subpopulations may be selected, which would lead to postzygotic isolation between isolated subpopulations. Our results indicate that sampling effects such as genetic drift or stochastic loss of Wolbachia produce postzygotic isolation in laboratory populations of spider mite.     

稻水象甲(Lissorhoptrus oryzophilus Kuschel)的卵子发生-飞行共轭

１９９４～１９９６年对双季稻区稻水象甲发生消长和迁飞动态的系统研究表明,稻水象甲的飞行肌和卵巢发育随生活史的季节性变化出现兴衰交替,从而迁入迁出稻田和越冬场所,表现出典型的卵子发生－飞行共轭：春季越冬代成虫从越冬场所迁入早稻田后飞行肌消解而卵巢发育,繁殖形成一代致害种群。夏季一代成虫生殖滞育,飞行肌和脂肪体发达,绝大部分个体迁出早稻田行夏蛰并越冬;少量落入秧田者和早稻收割时散落田内而晚稻插秧时尚未迁离的个体卵巢恢复发育,飞行肌消解而构成二代虫源。秋季二代成虫羽化后生殖滞育,迁飞上山越冬;１０月中旬后羽化的个体卵巢和飞行肌均不再发育而滞留田内外越冬。取食高质量食料时,一代夏蛰成虫飞行肌再生,新羽一代成虫则首先发育飞行肌,而后解除生殖滞育。     

A novel trade-off of insect diapause affecting a sequestered chemical defense

Diapause allows insects to temporally avoid conditions that are unfavorable for development and reproduction. However, diapause may incur a cost in the form of reduced metabolic energy reserves, reduced potential fecundity, and missed reproductive opportunities. This study investigated a hitherto ignored consequence of diapause: trade-offs involving sequestered chemical defense. We examined the aristolochic acid defenses of diapausing and non-diapausing pipevine swallowtail butterflies, Battus philenor. Pipevine swallowtail larvae acquire these chemical defenses from their host plants. Butterflies that emerge following pupal diapause have significantly less fat, a female fitness correlate, compared to those that do not diapause. However, butterflies emerging from diapaused pupae are more chemically defended compared to those that have not undergone diapause. Furthermore, non-diapausing butterflies are confronted with older, lower quality host plants on which to oviposit. Thus, a trade-off exists where butterflies may have greater energy reserves at the cost of less chemical defense and sub-optimal food resources for their larvae, or have substantially less energetic reserves with the benefit of greater chemical defense and plentiful larval food resources.     

Diapause termination, post-diapause development and reproduction in the beet webworm, Loxostege sticticalis (Lepidoptera: Pyralidae)

Effects of photoperiod and cold exposure on diapause termination, post-diapause development and reproduction in Loxostege sticticalis were examined. Larvae were reared at diapause inducing condition (22 °C, L:D 12:12) consistently or transferred to long day photoperiod (L:D 16:8) and darkness (L:D 0:24) respectively, after entering into diapause. Diapause was terminated in approximately 40% of the larvae after 36 days, and no significant differences were observed between photoperiods, suggesting larval diapause was terminated spontaneously without being induced by photoperiods. Cold exposure significantly hastened diapause termination. The diapause termination incidence increased significantly with peaks of 98% at both 5 °C and 0 °C exposure for 30 days, as compared to 42% in controls not exposed to cold, while the mortality and number of days required for diapause termination decreased dramatically. The optimal low temperature exposure periods under 5 °C or 0 °C were 20 days and 30 days, showing a higher termination incidence and shorter time for diapause termination. This suggests that the low temperatures in winter play an important role in diapause termination under natural conditions. The threshold temperatures for post-diapause development in prepupae and pupae were 9.13 °C and 10.60 °C respectively, with corresponding accumulations of 125 and 200 degree-days. Adults that experienced larval diapause significantly delayed their first oviposition, oviposition period was prolonged, and the lifetime number of eggs laid was decreased, however both males and females have significantly longer longevity. The field validation of diapause termination, the degree-days model, and the relationship between diapause and migration in L. sticticalis were also discussed.     

In vivo and in vitro effects of 20-hydroxyecdysone and methoprene on diapause maintenance and reproductive development in Musca autumnalis

Abstract. Face flies overwinter as adults in reproductive diapause. Administration of 20-hyroxyecdysone and/or methoprene induced reproductive development in diapausing flies which were maintained in a diapause-inducing environment. Hormone effects were additive and female flies were more sensitive than males. Release of vitellogenin from cultured fat body was stimulated by 20-hydroxyecdysone or methoprene. Transfer of flies from diapause to diapause-breaking environments induced some to break diapause, but this decreased with the time flies had been in a diapause-inducing environment. In contrast, topical application of methoprene to diapausing flies induced reproductive development irrespective of their ages even when they were kept in the diapause-inducing environment for 80 day degrees above a 12°C base temperature (14.5 days). Therefore diapause induction must depend on hormone levels less than some threshold level. The putative threshold varied according to diapause propensities of different genetic lines. Lines showing high frequencies of diapause required greater amounts of methoprene for reproductive development in diapause conditions than did lines showing low frequencies of diapause.     

Factors influencing the duration and termination of diapause in the Indian-meal moth, Plodia interpunctella

The influence of environmental factors on the duration of diapause in Plodia interpunctella larvae reared in short photoperiods at 20 or 25° C was examined, Diapause terminated most rapidly in long photoperiods at high temperatures. Pupation was more delayed, and mortality was higher, in darkness than in the presence of light. At 20° C, LD 16: 8 hastened diapause termination only slightly in unchilled samples. Chilling for 10 weeks at 10° C greatly reduced the duration of diapause at 20 or 25° C in constant darkness, and rendered LD 16:8 effective in terminating diapause at 20° C. In addition, the quite short duration of diapause under LD 16:8 at 25° C was further shortened by holding for 6–10 weeks at 10° C or below, or by holding in an outbuilding during winter. Holding diapausing larvae at 15 or 20° C proved less effective. Temperature rises from 20 to 25 or 30° C proved effective in terminating diapause. In one stock, the temperature at which diapause was induced influenced its subsequent duration. Lighting conditions during induction had less influence on duration than had temperature, and no difference occurred between pupation times of larvae reared at different population densities, Under all conditions tested, diapause lasted longer in a recently collected field stock than in a laboratory stock.     

Water relationships in the ectoparasitoid Nasonia vitripennis during larval diapause

Abstract. Final instar larvae of the ectoparasitoid Nasonia vitripennis overwinter in diapause within the puparium of the host fly. Wasp larvae that are in diapause have a different set of water relations than larvae that are not in diapause. Diapausing larvae contain less water, lose water at a lower rate, and have the capacity to absorb water vapour. Water conservation in larvae of N. vitripennis is also enhanced considerably by the puparium of its fly host.     

Trade-offs between storage and survival affect diapause timing in capital breeders

Many organisms spend the unfavourable part of the year, such as the winter season, in diapause or dormancy and reproduce in spring shortly after emergence. Reserves are acquired prior to diapause to cover metabolic costs and in some species also reproduction (capital breeding) directly after diapause. Storage is then a component of future reproduction, and capital breeders consequently pay a pre-breeding cost of reproduction as they risk dying while obtaining and carrying the reserves. How large should the reserves be, and to what extent should optimal storage, and thereby timing of diapause, depend on predation risk and reproductive strategy? We present a general and simplistic life history model of an arthropod (e.g. crustaceans or insects) that is exposed to background mortality risk when it accumulates reserves before diapause. The model optimizes diapause timing and resultant reserves for income, mixed and capital breeders, and predicts how mortality risk affects the degree of capital breeding. For income breeders, timing of diapause is insensitive to the risk while obtaining reserves as they, regardless of risk, acquire the minimum amount needed to survive the winter. For capital breeders, the higher the risk the earlier the diapause and less is consequently stored. Mixed breeders diapause late and store as much as pure capital breeders when exposed to low risk, but behave as income breeders and diapause early when mortality is high. Our model shows that the degree of capital breeding impacts phenology of diapause in a risk-dependent manner. This prediction should impact how diapause timing is thought of across a wide range of taxa, including the much studied marine copepods. Timing of diapause, including triggers and cues, can only be understood when the diversity of reproductive strategies and the adaptive value of storage is taken into account.     

Distribution of crustacean diapause: micro- and macroevolutionary pattern and process

Theoretical predictions for the relationships between duration of dormancy, reproductive life span, and dispersal ability developed for plants in temporally varying environments are applied here to crustaceans. Mathematical models suggest that diapause duration should negatively covary with adult life span, and that both diapause and life span should negatively covary with dispersal ability. A survey of 167 crustacean species from 20 orders and three classes confirms that species with prolonged diapause have short adult life spans and those with long adult lives either have diapause lasting less than a year, or do not diapause at all. Prolonged diapause is more common among small or inland water crustaceans than it is among large or marine species, whereas large or marine species have significantly longer adult life spans on average than do those that are small or from inland waters. A greater fraction of species in the Branchiopoda exhibit prolonged diapause than do members of the Maxillopoda which, in turn, are more likely to exhibit prolonged diapause than are the Malacostraca. A greater fraction of malacostracan species have adult life spans exceeding one year than do species in either the Branchiopoda or the Maxillopoda. Cladistic analysis shows that phylogenetic constraint is likely to be at least in part responsible for the expression of diapause among the Crustacea. We conclude that both natural selection and macroevolutionary pattern have influenced the distribution of diapause among modern crustaceans.     

Termination and induction of diapause in the gypsy moth larval parasitoid, Apanteles melanoscelus

Diapause in Apanteles melanoscelus can be terminated by exposure of the diapausing last instar larvae within their cocoons to 5°C for a period of 8 or more weeks. Photoperiod has no consistent influence upon diapause termination, but is of paramount importance for diapause induction. At less than 16 hr light per day virtually all larvae diapause, and at 18 hr and above very few larvae diapause. By exposing different larval stages to different photoperiods it was found that older larvae were most sensitive to the light-dark cycle. It was also noted that cocoons of diapausing larvae are larger than those of non-diapausing larvae.