Abstract: Diapause is a type of dormancy that requires preparation, typically precedes the onset of unfavorable conditions, and necessitates a period of arrest before development can proceed. Two ecologically important groups of copepods have incorporated diapausing stages into their life histories. In freshwater, estuarine, and coastal environments, species within the Centropagoidea superfamily can produce resting eggs containing embryos that may be quiescent, diapausing, or in some intermediate state. Resting eggs sink into the sediments, remain viable over months to years, and form a reservoir from which the planktonic population is reestablished. In coastal and oceanic environments, copepods within the Calanidae and Eucalanidae families can enter diapause during late juvenile (copepodid) or adult stages. These copepods accumulate large amounts of lipids before they migrate into deep water and diapause for several months. Through respiration, diapausing copepods may sequester more carbon in the deep ocean than a...

Abstract: Insects often overcome unfavorable seasons in a hormonally regulated state of diapause during which their activity ceases, development is arrested, metabolic rate is suppressed, and tolerance of environmental stress is bolstered. Diapausing insects pass through a stereotypic succession of eco-physiological phases termed “diapause development.” The phasing is varied in the literature, and the whole concept is sometimes criticized as being too artificial. Here we present the results of transcriptional profiling using custom microarrays representing 1,042 genes in the drosophilid fly, Chymomyza costata. Fully grown, third-instar larvae programmed for diapause by a photoperiodic (short-day) signal were assayed as they traversed the diapause developmental program. When analyzing the gradual dynamics in the transcriptomic profile, we could readily distinguish distinct diapause developmental phases associated with induction/initiation, maintenance, cold acclimation, and termination by cold or by photoperiodic signal. Accordingly, each phase is characterized by a specific pattern of gene expression, supporting the physiological relevance of the concept of diapause phasing. Further, we have dissected in greater detail the changes in transcript levels of elements of several signaling pathways considered critical for diapause regulation. The phase of diapause termination is associated with enhanced transcript levels in several positive elements stimulating direct development (the 20-hydroxyecdysone pathway: Ecr, Shd, Broad; the Wnt pathway: basket, c-jun) that are countered by up-regulation in some negative elements (the insulin-signaling pathway: Ilp8, PI3k, Akt; the target of rapamycin pathway: Tsc2 and 4EBP; the Wnt pathway: shaggy). We speculate such up-regulations may represent the early steps linked to termination of diapause programming.

Abstract: Diapause, a strategy to endure unfavourable conditions (e.g. cold winters) is commonly found in ectothermic organisms and is characterized by an arrest of development and reproduction, a reduction of metabolic rate, and an increased resistance to adversity. Diapause, in addition to adaptations for surviving low winter temperatures, significantly influences phenology, voltinism and ultimately population growth. We review the literature on diapause and overwintering behaviour of two bark beetle species that affect spruce-dominated forests in the northern hemisphere, and describe and compare how these strategies can influence population dynamics. The European spruce bark beetle Ips typographus (L.) (Coleoptera, Curculionidae) is the most important forest pest of Norway spruce in Europe. It enters an adult reproductive diapause that might be either facultative or obligate. Obligate diapausing beetles are considered strictly univoltine, entering this dormancy type regardless of environmental cues. Facultative diapausing individuals enter diapause induced by photoperiod, modified by temperature, thus being potentially multivoltine. The spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) infests all spruce species in its natural range in North America. A facultative prepupal diapause is averted by relatively warm temperatures, resulting in a univoltine life cycle, whereas cool temperatures induce prepupal diapause leading to a semivoltine cycle. An adult obligate diapause in D. rufipennis could limit bi- or multivoltinism. We discuss and compare the influence of diapause and overwinter survival on voltinism and population dynamics of these two species in a changing climate and provide an outlook on future research.

Abstract: Diapause, also known as dormancy, is a state of arrested development that allows insects to survive unfavorable environmental conditions. Diapause-destined insects store large amounts of fat when preparing for diapause. However, the extent to which these accumulated fat reserves influence diapause remains unclear. To address this question, we investigated the function of fatty acid synthase (FAS), which plays a central role in lipid synthesis, in stress tolerance, the duration of diapause preparation, and whether insects enter diapause or not. In diapause-destined adult female cabbage beetles, Colaphellus bowringi, FAS2 was more highly expressed than FAS1 at the peak stage of diapause preparation. FAS2 knockdown suppressed lipid accumulation and subsequently affected stress tolerance genes expression and water content. However, silencing FAS2 had no significant effects on the duration of diapause preparation or the incidence of diapause. FAS2 transcription was suppressed by juvenile hormone (JH) and the JH receptor methoprene-tolerant (Met). These results suggest that the absence of JH-Met induces FAS2 expression, thereby promoting lipid storage in diapause-destined female beetles. These results demonstrate that fat reserves regulate stress tolerance genes expression and water content, but have no significant effect on the duration of diapause preparation or the incidence of diapause.

Abstract: Speciation with gene flow may require adaptive divergence of multiple traits to generate strong ecologically based reproductive isolation. Extensive negative pleiotropy or physical linkage of genes in the wrong phase affecting these diverging traits may therefore hinder speciation, while genetic independence or “modularity” among phenotypic traits may reduce constraints and facilitate divergence. Here, we test whether the genetics underlying two components of diapause life history, initial diapause intensity and diapause termination timing, constrain differentiation between sympatric hawthorn and apple-infesting host races of the fly Rhagoletis pomonella through analysis of 10,256 SNPs measured via genotyping-by-sequencing (GBS). Loci genetically associated with diapause termination timing were mainly observed for SNPs mapping to chromosomes 1–3 in the genome, most notably for SNPs displaying higher levels of linkage disequilibrium (LD), likely due to inversions. In contrast, selection on initial diapause intensity affected loci on all five major chromosomes of the genome, specifically those showing low levels of LD. This lack of overlap in genetically associated loci suggests that the two diapause phenotypes are largely modular. On chromosome 2, however, intermediate level LD loci and a subgroup of high LD loci displayed significant negative relationships between initial diapause intensity and diapause termination time. These gene regions on chromosome 2 therefore affected both traits, while most regions were largely independent. Moreover, loci associated with both measured traits also tended to exhibit highly divergent allele frequencies between the host races. Thus, the presence of nonoverlapping genetic modules likely facilitates simultaneous, adaptive divergence for the measured life-history components.

Abstract: In temperate insects, winters are typically endured by entering diapause, which comprises a deep resting stage. Correct timing of diapause termination is vital for synchronization of emergence with conspecifics and for mobilizing resources when conditions for growth and reproduction become favourable. Although critical to survival, the intrinsic and extrinsic drivers of diapause termination timing are poorly understood. In the present study, we investigate diapause development under a range of durations (10–24 weeks) spent at different temperatures (−2 to 10 °C) in the pupal diapausing butterfly Pieris napi Linnaeus (Lepidoptera:Pieridae). We determine: (i) the maximum cold temperature for diapause development; (ii) if pupae in diapause count cold days or cold sums; and (iii) whether diapause termination is distinct or gradual. The results indicate large and idiosyncratic effects of high and low nonlethal temperatures on diapause development in P. napi. Although all temperatures tested lead to diapause termination, a thermal optimum between 2 and 4 °C is observed. Lower temperatures lead to decreased eclosion propensity, whereas higher temperatures slow down development and increase emergence desynchronization. These data suggest that, rather than a simple cold-summing process with a distinct diapause termination point, there are trade-offs between time and temperature at the low and high end of the thermal range, resulting in a nonlinear thermal landscape showing a ridge of increasing eclosion propensity at moderate temperatures. The present study suggests that the effects of temperature on diapause development should be included in projections on post-winter phenology models of insects, including pest species.

Abstract: Diapause is an endogenously regulated dormant state that allows insects and other animals to survive seasons of inimical conditions Diapause is characterized by altered gene expression profiles, which mediate developmental arrest, metabolic depression, increased stress resistance, and other physiological and biochemical changes associated with this dormant state In this chapter, I explore epigenetic processes that may regulate gene expression and contribute to the diapause phenotype Only a few studies have explored the influence of DNA methylation, histone modification, and small RNA silencing on diapause However, there is evidence that histone methylation and histone acetylation influence diapause in a wide range of species In addition, microRNAs, siRNAs, and piRNAs appear to regulate gene expression before, during, and after diapause Together, the accumulating evidence supports the inclusion of multiple epigenetic pathways in the “toolbox” of processes that regulate dormant states in insects and other animals

Abstract: Diapause is a programmed dormancy that allows organisms to tolerate predictable periods of unfavourable conditions by temporarily halting development and reducing metabolism. Diapause is widespread amongst insects and is crucial for allowing organisms to coordinate their growth and reproduction with favourable environmental conditions. Although the adaptive significance of diapause is well understood, the molecular mechanisms underpinning diapause remain unresolved. We performed high-throughput sequencing to investigate the role of microRNAs (miRNAs) in the diapause of the Asian tiger mosquito, Aedes albopictus. We first investigated miRNAs in diapause induction by characterizing maternally provisioned miRNAs in mature oocytes of Ae. albopictus under diapause-inducing and diapause-averting conditions. Second, we investigated miRNAs in diapause maintenance by characterizing miRNAs in diapause and nondiapause pharate larvae. We identified 162 miRNAs, 152 previously known and 10 putatively novel. We identified no differentially abundant miRNAs in mature oocytes and seven differentially abundant miRNAs in pharate larvae. The predicted targets of differentially abundant miRNAs include genes affecting several processes related to diapause maintenance including ecdysone regulation, immune response, lipid metabolism and regulation of development. Our results suggest that Ae. albopictus does not maternally provision a unique set of miRNAs during diapause induction but miRNAs are a component of diapause maintenance in this species.

Abstract: Temperature-based degree-day models describe insect seasonality and to predict key phenological events. We expand on the use of a temperature-based process defining timing of reproduction through the incorporation of female reproductive physiology for the invasive pentatomid species Halyomorpha halys, the brown marmorated stink bug. A five-stage ranking system based on ovary development was able to distinguish between the reproductive statuses of field-collected females. Application of this ranking method described aspects of H. halys' seasonality, overwintering biology, and phenology across geographic locations. Female H. halys were collected in the US from NJ, WV, NC, OR, and two sites in PA in 2006-2008 (Allentown, PA only) and 2012-2014. Results identify that H. halys enters reproductive diapause in temperate locations in the fall and that a delay occurs in developmental maturity after diapause termination in the spring. Modification of the Snyder method to identify biofix determined 12.7-hr photoperiod as the best fit to define initiation of reproduction in the spring. Applying the biofix, we demonstrated significant differences between locations for the rate at which the overwintering generation transition into reproductive status and the factors contributing to this difference require further study. For example, after including abiotic variables influencing development such as temperature and photoperiod (critical diapause cue), reproduction occurred earlier in OR and for an extended period in NJ. This data describe a method to investigate insect seasonality by incorporating physiological development across multiple regions that can clarify phenology for insects with overlapping generations.

Abstract: Many insects survive seasonal adversities during diapause, a form of programmed developmental and metabolic arrest. Photoperiodically regulated entry into diapause allows multivoltine insect species to optimize the number of generations. The molecular mechanism of the photoperiodic timer is unknown in insects. In the present study, we take advantage of the robust reproductive diapause response in the linden bug Pyrrhocoris apterus and explore the fifth-instar nymphal stage, which is the most photoperiod-sensitive stage. The nymphs display daily changes in locomotor activity during short days; this differs from the activity observed during long days. We find evidence of cyclical expression of the circadian clock genes, per and cyc, in nymphal heads; in addition, per expression is also photoperiod-dependent. The RNA interference-mediated knockdown of the two circadian clock genes, Clk and cyc, during the nymphal stage results in reproductive arrest in adult females. Furthermore, Clk and cyc knockdown induces the expression of the storage protein hexamerin in the fat body, whereas the expression of vitellogenin diminishes. Taken together, these data support the involvement of circadian clock genes in photoperiodic timer and/or diapause induction.

Abstract: Khapra beetle, Trogoderma granarium Everts (Coleoptera: Dermestidae), is a pest of stored grain in Africa, Asia, and Europe. It is a quarantine insect for much of the rest of the world. Control of T. granarium can be achieved with methyl bromide, but this fumigant is an ozone‐depleting substance and is being phased out worldwide. Thus, there is an urgent need to find new methods of control, including the use of low temperatures. Here, we assess the effects of diapause and cold acclimation on the cold tolerance of T. granarium. The percentage of larvae in diapause increased with larval density, reaching 57.3% when reared at a density of 73 larvae g−1 diet. The cold tolerance of T. granarium was assessed by the supercooling points (SCPs) of various life stages. The SCP of non‐acclimated insects ranged from −26.2 ± 0.2 °C (mean ± SEM) for eggs to −14.4 ± 0.4 °C for larvae. The lowest SCP for larvae, −24.3 ± 0.3 °C, was obtained for diapausing‐acclimated larvae. Based on mean LT50 values, the most cold‐tolerant stage at −10 °C was the diapausing‐acclimated larvae (87 days) followed by non‐diapausing‐acclimated larvae (51 days), diapausing non‐acclimated larvae (19 days), adults (4 days), non‐diapausing non‐acclimated larvae (2 days), pupae (0.4 days), and eggs (0.2 days). The estimated times to obtain 99.9968% mortality (Probit 9) for diapausing‐acclimated larvae are 999, 442, 347, 84, and 15 days at 0, −5, −10, −15, and −20 °C, respectively. Probit 9 is an estimated value used by quarantine experts to estimate conditions that are required to kill all insects. In light of the long exposure time needed to control T. granarium even at −20 °C, cooling to below −27 °C (i.e., below the SCP of eggs) will quickly kill all life stages and may be the best way to control this insect with low temperatures.

Abstract: Prior to entering diapause, insects must prepare themselves physiologically to withstand the stresses of arresting their development for a lengthy period. While studies describing the biochemical and cellular milieu of the maintenance phase of diapause are accumulating, few studies have taken an ‘omics’ approach to describing molecular events during the diapause preparatory phase. We used isobaric tags and mass spectrometry (iTRAQ) to quantitatively compare the expression profiles of proteins identified during the onset of diapause preparation phase in the heads of adult female cabbage beetles, Colaphellus bowringi. A total of 3,175 proteins were identified, 297 of which were differentially expressed between diapause-destined and non-diapause-destined female adults and could therefore be involved in diapause preparation in this species. Comparison of identified proteins with protein function databases shows that many of these differentially expressed proteins enhanced in diapause destined beetles are involved in energy production and conversion, carbohydrate metabolism and transport, and lipid metabolism. Further hand annotation of differentially abundant peptides nominates several associated with stress hardiness, including HSPs and antioxidants, as well as neural development. In contrast, non-diapause destined beetles show substantial increases in cuticle proteins, suggesting additional post-emergence growth. Using RNA interference to silence a fatty acid-binding protein that was highly abundant in the head of diapause-destined females prevented the accumulation of lipids in the fat body, a common product of diapause preparation in this species and others. Surprisingly, RNAi against the fatty acid-binding protein also affected the transcript abundance of several heat shock proteins. These results suggest that the identified differentially expressed proteins that play vital roles in lipid metabolism may also contribute somehow to enhanced hardiness to environmental stress that is characteristic of diapause.

Abstract: Killifishes survive and persist in extreme environments by exploiting both aquatic and terrestrial habitats for egg deposition, and by adjusting the length of development to match availability of water to support larval growth and maturation. Annual killifishes persist in ephemeral bodies of water through the production of drought-tolerant embryos. Survival of the environmental stresses associated with their highly variable and seasonal habitat is supported by their ability to enter into at least two states of metabolic and developmental dormancy, diapause or quiescence. There are three stages of diapause in annual killifishes, one occurring prior to gastrulation, one about midway through development, and one in late pre-hatching embryos. Quiescence may occur at any developmental stage. In addition, delayed hatching is known to occur in close relatives of the annual killifishes, and may be superficially confused with pre-hatching diapause. These types of developmental delay are induced by different cues and serve different purposes in the life history of the species. Thus, it is likely that the molecular mechanisms that induce dormancy and support survival are unique in each case. It is imperative that we properly define these forms of developmental dormancy in our studies in order to put our results into the proper ecological and evolutionary context. Here the unique characteristics of these distinct categories of developmental delay are reviewed. Developmental Dynamics 246:858-866, 2017. © 2017 The Authors Developmental Dynamics published by Wiley Periodicals, Inc. on behalf of American Association of Anatomists.

Abstract: Temperature is both a selective pressure and a modulator of the diapause expression in insects from temperate regions. Thus, with climate warming, an alteration of the response to seasonal changes is expected, either through genetic adaptations to novel climatic conditions or phenotypic plasticity. Since the 1980s in western France, the winter guild of aphid parasitoids (Hymenoptera: Braconidae) in cereal fields has been made up of two species: Aphidius rhopalosiphi and Aphidius matricariae. The recent activity of two other species, Aphidius avenae and Aphidius ervi, during the winter months suggests that a modification of aphid parasitoid overwintering strategies has taken place within the guild. In this study, we first performed a field survey in the winter of 2014/15 to assess levels of parasitoid diapause incidence in agrosystems. Then, we compared the capacity of the four parasitoid species to enter winter diapause under nine different photoperiods and temperature conditions in the laboratory. As predicted, historically winter-active species (A. rhopalosiphi and A. matricariae) never entered diapause, whereas the species more recently active during winter (A. avenae and A. ervi) did enter diapause but at a low proportion (maximum of 13.4 and 11.2%, respectively). These results suggest rapid shifts over the last three decades in the overwintering strategies of aphid parasitoids in Western France, probably due to climate warming. This implies that diapause can be replaced by active adult overwintering, with potential consequences for species interactions, insect community composition, ecosystem functioning, and natural pest control.

Abstract: Many temperate insects survive harsh environmental conditions, such as winter, by entering a state of developmental arrest. This diapause state is predominantly induced by photoperiod. The photoper ...

Abstract: Diapause is a deep resting stage facilitating temporal avoidance of unfavourable environmental conditions, and is used by many insects to adapt their life cycle to seasonal variation. Although cons ...

Abstract: Embryonic diapause is a common reproductive strategy amongst mammals, requiring an intimate cross-talk between the endometrium and the blastocyst. To date, the precise molecular signals responsible are unknown in the mouse or any other mammal. Previous studies in the mink implicate polyamines as major regulators of the control of diapause. In the mouse, inhibiting the rate-limiting enzyme of polyamine synthesis, ornithine decarboxylase (ODC1) during early pregnancy largely prevents implantation, but the fate of the nonimplanted embryos is unknown. To determine whether polyamines control mouse embryonic diapause, we treated pregnant mice with an ODC1 inhibitor from d3.5 to d6.5 postcoitum. At d7.5, 72% of females had no signs of implantation whilst the remaining females exhibited disrupted placental formation and degenerate embryos. In the females with no implantation, we obtained viable blastocysts that had attenuated cell proliferation, indicating a state of diapause. When cultured in vitro, these exhibited trophoblast outgrowth, indicative of reactivation of embryogenesis. In contrast, direct culture of d3.5 blastocysts with an ODC1 inhibitor failed to cause entry into diapause. Examination of the polyamine pathway enzymes and a number of implantation factors indicated inhibition of ODC1 resulted in a uterine phenotype that resembled diapause, with some compensatory increases in crucial genes. Thus, we conclude that an absence or paucity of polyamines induces the uterine quiescence that causes entry of the blastocyst into embryonic diapause.

Abstract: During unfavourable conditions, many arthropods have the ability to enter into diapause and synchronize their development and reproduction to seasonal patterns. Diapause or winter hibernation in insects and mites is set off by a number of cues, with photoperiod being the most well-defined and strongest signal. This review focuses on the current knowledge of '-omics' data and the genetics of diapause in the two-spotted spider mite Tetranychus urticae, a member of the family Tetranychidae (Arthropoda: Chelicerata: Arachnida: Acari). This species is a serious polyphagous pest and females undergo a reproductive facultative diapause when immature stages are exposed to long nights. Winter hibernation induces different physiological processes characterized by a metabolic suppression, different energy use, increased stress tolerance and the production of cryoprotectants, all initiated by a complex signal transduction pathway. Keto-carotenoids are known to cause the deeply orange colour typical for diapausing females. Furthermore, research with colour mutants of T. urticae has shown the need for carotenoids with respect to the induction of diapause, even though the molecular-genetic mechanisms underlying these colour phenotypes are still unknown. In addition, marked latitudinal variation in diapause incidence among populations has been observed in nature, with modes of inheritance ranging from recessive to dominant, as well as monogenic to polygenic. We end by highlighting the emerging opportunities for functional studies that aim to unravel the complex factors underlying diapause in spider mites.

Abstract: We examined age-related biochemical and histological changes in the fat bodies and hemolymph of Osmia rufa males and females. We analysed solitary bees during diapause, in October and in April; as well as the flying insects following diapause, in May and June. The trophocyte sizes, as well as the numbers of lipid droplets were the greatest at the beginning of diapause. Subsequently, they decreased along with age. Triglyceride and glucose concentrations systematically decreased in fat body cells but increased in the hemolymph from October to June. Concentrations/activities of (enzymatic and non-enzymatic) antioxidant and proteolytic systems, as well as phenoloxidase, aspartate aminotransferase, alanine aminotransferase and alkaline phosphatase levels were constant during the diapause, usually lower in the males than the females. Prior to the diapause/overwintering, the concentrations/activities of all the compounds were higher in the fat bodies than in the hemolymph. Later in the spring and in the summer, they increased in the hemolymph and on the body surfaces, while decreasing in the fat bodies. The global DNA methylation levels increased with age. Higher levels were always observed in the males than in the females. The study will promote better understanding of bee evolution and will be useful for the protection and management of solitary bees, with benefits to the environment and agriculture.

Abstract: Diapause, which occurs through the production of dormant eggs, is a strategy used by some zooplankton to avoid winter months of persistent low temperatures and low food availability. However, reports of active zooplankton under the ice indicate that other strategies also exist. This study was aimed at evaluating whether the composition of storage lipids in the fall differs between diapausing and active overwintering Daphnia. We assessed the quantity of storage lipids and fatty acid (FA) composition of Daphnia species, along with FA content of seston, in six boreal, alpine and subarctic lakes at the onset of winter, and evaluated the association between storage lipids and Daphnia overwintering strategy. We found that active overwintering Daphnia had >55% body fat and the highest FA concentrations. Polyunsaturated FA, especially stearidonic acid (18:4n-3; SDA) and high ratios of n-3:n-6, were preferentially retained to a greater extent in active overwintering Daphnia than in those that entered diapause. Daphnia FA composition was independent of that of the seston diet, indicating that Daphnia adjusted their storage lipids according to the physiological requirements of a given overwintering strategy. The occurrence of an active overwintering strategy has consequences for zooplankton community structure, and can have important implications for the transfer of high-quality energy at higher trophic levels.

Abstract: This paper reviews our very limited knowledge about resting-egg hatching and early population development in planktonic rotifers. Hatching of stem females from resting eggs may occur soon after resting eggs are produced, but perhaps usually it is delayed by a minimum obligate diapause, by a requirement for seasonal temperature changes, or by sinking to sediment environments that prevent hatching. In deep-water sediments, hatching probably is inhibited by low oxygen, darkness, or low temperature, so that eggs likely hatch following resuspension during water-column turnover. Populations should develop primarily by female parthenogenesis and have relatively low clonal diversity. By contrast, in shallow-water sediments, eggs are more likely to experience conditions conducive to hatching and to be resuspended into the water column. Populations may develop by massive emergence of stem females as well as by female parthenogenesis and thus have a very high clonal diversity. Stem females of some species are particularly fit for colonization of new habitat. First, compared to females hatched from parthenogenetic eggs, they can have a greater lipid reserve that enhances survival and reproduction. Second, amictic stem females can contain a transmissible factor that inhibits sexual reproduction and diapause for several to many successive generations, thus promoting rapid reproduction via female parthenogenesis.

Abstract: Diapause, a programmed developmental arrest, is common in insects, enabling them to survive adverse seasons. It is well established that pupal diapause is regulated by ecdysteroids secreted by the prothoracic glands (PGs), with cessation of ecdysteroid secretion after pupal ecdysis leading to pupal diapause. A major factor regulating the gland activity is prothoracicotropic hormone (PTTH) secreted from the brain. In our previous study, we demonstrated that the cessation of PTTH release after pupal ecdysis resulted in the inactivation of the PGs, leading to pupal diapause in the cabbage army moth Mamestra brassicae. Here we show that a neuropeptide myosuppressin also contributes to the inactivation of PGs at the initiation of diapause. Myosuppressin suppresses PTTH-stimulated activation of the PGs in vitro. Concentrations of myosuppressin in the hemolymph after pupal ecdysis are higher in diapause pupae than in nondiapause pupae.

Abstract: Insects living in the temperate climate include summer diapause, or aestivation, in their seasonal cycle to solve various problems related to adaptation to unfavorable seasons. Unlike winter diapause, summer diapause occurs in summer and is usually terminated in autumn when active feeding, development, and/or reproduction are restored. Typically, high temperature and long day induce summer diapause and then maintain it, whereas short day and low temperature prevent induction of this diapause or terminate it. The summer diapause syndrome is basically similar to that of winter diapause; it includes prior development of large fat body, decreased level of metabolism, increased general resistance to unfavorable abiotic and biotic conditions, etc. Inhibition of morphogenesis and gametogenesis is under the control of the endocrine system. The onset of summer diapause is often accompanied by migrations to varying, sometimes significant distances to the sites of aestivation. The selective factors responsible for evolution of summer diapause vary between insect species. Climatic factors and, consequently, availability and abundance of food, as well as pressure of predators and parasites are likely to be the main factors that stimulate its occurrence. In some species, prolonged diapause begins in spring or early summer and ceases only after over-wintering. When studied in detail, such prolonged diapause often turns out to be a sequence of two independent diapauses, summer and winter ones, occurring in succession without detectable external changes.

Abstract: The lifespan of Drosophila melanogaster can be extended substantially by inducing reproductive dormancy (also known as diapause) by lowered temperature and short days. This increase of longevity is accompanied by lowered metabolism and increased stress tolerance. We ask here whether behavioral senescence is ameliorated during adult dormancy. To study this we kept flies for seven or more weeks in normal rearing conditions or in diapause conditions and compared to one-week-old flies in different behavioral assays of sleep, negative geotaxis and exploratory walking. We found that the senescence of geotaxis and locomotor behavior seen under normal rearing conditions was negligible in flies kept in dormancy. The normal senescence of rhythmic activity and sleep patterns during the daytime was also reduced by adult dormancy. Investigating the morphology of specific neuromuscular junctions we found that changes normally seen with aging do not take place in dormant flies. To monitor age-associated changes in neuronal circuits regulating activity rhythms, sleep and walking behavior we applied antisera to tyrosine hydroxylase, serotonin and several neuropeptides to examine changes in expression levels and neuron morphology. In most neuron types the levels of stored neuromodulators decreased during normal aging, but not in diapause treated flies. No signs of neurodegeneration were seen in either condition. Our data suggest that age-related changes in motor neurons could be the cause of part of the behavioral senescence and that this is ameliorated by reproductive diapause. Earlier studies established a link between age-associated decreases in neuromodulator levels and behavioral decline that could be rescued by overexpression of neuromodulator. Thus, it is highly likely that the retained levels of neuromodulators in dormant flies alleviate behavioral senescence.

Abstract: To cope with temporal and spatial heterogeneity of habitats, herbivorous insects in the temperate zone usually enter diapause that facilitates synchronization of their life cycle with specific stages of host plants, such as fruit ripening. In the present study, we address those factors regulating dormancy responses as part of a ‘longer strategy’ to persist and thrive in temperate environments, focusing on Rhagoletis cerasi, a univoltine, oligophagous species, which overwinters as pupae and emerges when host fruits are available for oviposition at local scale. To ensure population survival and reproduction at habitats with ecological heterogeneity, R. cerasi has evolved a sophisticated diapause strategy based on a combination of local adaptation and diversified bet-hedging strategies. Diapause duration is determined both by (i) the adaptive response to local host fruit phenology patterns (annual diapause) and (ii) the plastic responses to unpredictable inter-annual (temporal) climatic variability that drives a proportion of the populations to extend dormancy by entering a second, successive, facultative cycle of prolonged diapause as part of a bet-hedging strategy. Besides the dormant periods, post-diapause development (which varies among populations) exerts ‘fine tune’ adjustments that assure synchronization and may correct possible errors. Adults emerging from pupae with prolonged diapause are larger in body size compared with counterparts emerging during the first year of diapause. However, female fecundity rates are reduced, followed by an extended post-oviposition period, whereas adult longevity remains unaffected. Overall, it appears that R. cerasi populations are adapted to ecological conditions of local habitats and respond plastically to unpredictable environmental (climatic) conditions.