Variation In Diapause Research Articles

Partial reuse of circadian clock genes along parallel clines of diapause in two moth species.

Understanding the molecular basis of repeated evolution improves our ability to predict evolution across the tree of life. Only since the last decade has high-throughput sequencing enabled comparative genome scans to thoroughly examine the repeatability of genetic changes driving repeated phenotypic evolution. The Asian corn borer (ACB), Ostrinia furnacalis (Guenée), and the European corn borer (ECB), Ostrinia nubilalis (Hübner), are two closely related moths displaying repeatable phenological adaptation to a wide range of climates on two separate continents, largely manifesting as changes in the timing of diapause induction and termination across latitude. Candidate genes underlying diapause variation in North American ECB have been previously identified. Here, we sampled seven ACB populations across 23 degrees of latitude in China to elucidate the genetic basis of diapause variation and evolutionary mechanisms driving parallel clinal responses in the two species. Using pooled whole-genome sequencing (Pool-seq) data, population genomic analyses revealed hundreds of single nucleotide polymorphisms (SNP) whose allele frequencies covaried with mean diapause phenotypes along the cline. Genes involved in circadian rhythm were over-represented among candidate genes with strong signatures of spatially varying selection. Only one of two circadian clock genes associated with diapause evolution in ECB showed evidence of reuse in ACB (period [per]), but per alleles were not shared between species nor with their outgroup, implicating independent mutational paths. Nonetheless, evidence of adaptive introgression was discovered at putative diapause loci located elsewhere in the genome, suggesting that de novo mutations and introgression might both underlie the repeated phenological evolution.

Unique genetic signatures of local adaptation over space and time for diapause, an ecologically relevant complex trait, in Drosophila melanogaster.

Organisms living in seasonally variable environments utilize cues such as light and temperature to induce plastic responses, enabling them to exploit favorable seasons and avoid unfavorable ones. Local adapation can result in variation in seasonal responses, but the genetic basis and evolutionary history of this variation remains elusive. Many insects, including Drosophila melanogaster, are able to undergo an arrest of reproductive development (diapause) in response to unfavorable conditions. In D. melanogaster, the ability to diapause is more common in high latitude populations, where flies endure harsher winters, and in the spring, reflecting differential survivorship of overwintering populations. Using a novel hybrid swarm-based genome wide association study, we examined the genetic basis and evolutionary history of ovarian diapause. We exposed outbred females to different temperatures and day lengths, characterized ovarian development for over 2800 flies, and reconstructed their complete, phased genomes. We found that diapause, scored at two different developmental cutoffs, has modest heritability, and we identified hundreds of SNPs associated with each of the two phenotypes. Alleles associated with one of the diapause phenotypes tend to be more common at higher latitudes, but these alleles do not show predictable seasonal variation. The collective signal of many small-effect, clinally varying SNPs can plausibly explain latitudinal variation in diapause seen in North America. Alleles associated with diapause are segregating in Zambia, suggesting that variation in diapause relies on ancestral polymorphisms, and both pro- and anti-diapause alleles have experienced selection in North America. Finally, we utilized outdoor mesocosms to track diapause under natural conditions. We found that hybrid swarms reared outdoors evolved increased propensity for diapause in late fall, whereas indoor control populations experienced no such change. Our results indicate that diapause is a complex, quantitative trait with different evolutionary patterns across time and space.

Swede midge (Diptera: Cecidomyiidae) diapause initiation under stable conditions: not a family affair

AbstractLarvae of the swede midge,Contarinia nasturtii(Kieffer) (Diptera: Cecidomyiidae), rely on environmental cues to initiate the diapause programme. This facultative diapause strategy, which provides opportunities for both bet hedging and multivoltinism, likely contributed to the successful colonisation and spread of this invasive pest in North America. Population diapause incidence is variable even under relatively stable conditions (especially at the critical day length), yet the mechanisms influencing the developmental trajectory of individuals under such conditions are unknown. Here we approach the non-environmental (i.e., heritable) controls of this variation by comparing diapause frequencies within and among full-sibling swede midge families reared in one of two stable environments. Under warm, long-day conditions > 99% of swede midges pupated directly, while 86% of swede midges entered diapause under cool, short-day conditions. In the latter condition, most families exhibited mixed developmental trajectories (ranging from 36% to 96% diapause). This developmental variation among siblings indicates that the diapause induction threshold does not follow simple Mendelian inheritance with complete dominance for a particular allele, but may follow incomplete dominance or a more complex heredity. Alternatively, within-family diapause variation may result from maternal bet hedging or factors such as maternal age or larval nutrition.

Geographic Variation of Diapause Induction Rates in Trichogramma drendrolimi (Hymenoptera: Trichogrammatidae) in China.

Diapause variation is a key factor affecting the development and success of parasitoids employed as biological control agents. Trichogramma dendrolimi Matsumura is mass-reared for biocontrol purposes in several countries around the world. Here we investigated the rate of diapause induction in three populations of T. dendrolimi from different regions in China: Heilongjiang (44° 55' N, 128° 26' E; HLJ), Liaoning (40° 18' N, 123° 22' E; LN), and Jiangsu province (32° 30' N, 120° 09' E; JS). We measured the rates of diapause induction for populations from each region when exposed to temperatures of 8, 10, 12, and 14 °C, for periods ranging from 5 to 35 d. Results showed that both the temperature and exposure duration influenced diapause induction in the three populations of T. dendrolimi. The HLJ and LN populations showed the highest percentage of diapause under 12 and 10 °C, regardless of time period tested. After 20 d at 12 and 10 °C, 97.42% and 95.94% of individuals of HLJ and LN, respectively, entered diapause. After treatment for 20-35 d under 8 °C, or 15-25 d under 10 °C, the diapause rate of LN was higher than those of the other two populations. In contrast, the higher temperatures (12-14 °C) induced diapause rapidly and steadily in HLJ. The diapause rate of JS was low for all temperatures and time periods. After 30 d at 12 °C, the JS population reached its highest diapause rate (29.56%). Our results showed significant variance in diapause rates among geographically distinct populations of T. dendrolimi in response to various abiotic conditions.

Divergence of the diapause transcriptome in apple maggot flies: winter regulation and post-winter transcriptional repression.

The duration of dormancy regulates seasonal timing in many organisms and may be modulated by day length and temperature. Though photoperiodic modulation has been well studied, temperature modulation of dormancy has received less attention. Here, we leverage genetic variation in diapause in the apple maggot fly, Rhagoletis pomonella, to test whether gene expression during winter or following spring warming regulates diapause duration. We used RNAseq to compare transcript abundance during and after simulated winter between an apple-infesting population and a hawthorn-infesting population where the apple population ends pupal diapause earlier than the hawthorn-infesting population. Marked differences in transcription between the two populations during winter suggests that the 'early' apple population is developmentally advanced compared with the 'late' hawthorn population prior to spring warming, with transcripts participating in growth and developmental processes relatively up-regulated in apple pupae during the winter cold period. Thus, regulatory differences during winter ultimately drive phenological differences that manifest themselves in the following summer. Expression and polymorphism analysis identify candidate genes in the Wnt and insulin signaling pathways that contribute to population differences in seasonality. Both populations remained in diapause and displayed a pattern of up- and then down-regulation (or vice versa) of growth-related transcripts following warming, consistent with transcriptional repression. The ability to repress growth stimulated by permissive temperatures is likely critical to avoid mismatched phenology and excessive metabolic demand. Compared with diapause studies in other insects, our results suggest some overlap in candidate genes/pathways, though the timing and direction of changes in transcription are likely species specific.

Seasonal and geographical variation in diapause and cold hardiness of the Asian corn borer, Ostrinia furnacalis.

Asian corn borer, Ostrinia furnacalis (Guenée), is a key corn pest in the Asian-Western Pacific countries. It overwinters as full-grown larvae in plant stalks or in a spun-silk covering located in the plant debris in the temperate regions of China. Supercooling point (SCP) and survival rate after low sub-zero temperature treatment were assessed for field-collected populations in the laboratory using a cool bath with a 1°C/min cooling rate until -40°C. Mean SCPs were varied among geographical populations, with a significant decline from -22.7°C of Haikou, the multivoltine tropical population in the south, to -28.5°C of Gongzhuling, the univoltine temperate population in the northeast of China. In addition, there was more than 1°C difference in SCP between Gongzhuling univoltine and bivoltine populations that were from the same geographic origin. Mean SCPs of the Guangzhou population fluctuated over the year, with significantly lower SCPs in winter than in other seasons, which correlated with a significantly higher proportion of diapausing larvae in winter than in other seasons. Over 41% of overwintering larvae from the northeast population could withstand to be supercooled for a few minutes to the low sub-zero temperature of -40°C, but only 6.7% of their southern counterparts did so. The findings from this study suggest that O. furnacalis mostly takes advantage of freeze avoidance as diapausing larvae for overwintering in the southern region, whereas it exhibits freeze tolerance in diapause in the northeastern region.

Months of Asynchrony in Offspring Production but Synchronous Adult Emergence: The Role of Diapause in an Ectoparasite's Life Cycle

Off-host stages of temperate zone ectoparasites must overcome two challenges: coping with unfavorable seasons and synchronizing their life cycles with host availability. In general, little is known about the seasonal cycles of insect ectoparasites of warm-blooded animals. The current study investigates the unusual phenology of a viviparous hippoboscid fly, the deer ked (Lipoptena cervi L.), that parasitizes boreal cervids. Despite months of asynchrony in offspring production, the adults emerge synchronously in mid-August across the northern boreal zone. We examined the role of diapause variation in the synchronization of life cycles by testing adult emergence success and time in relation to offspring birth month (October to April) and with respect to chilling time and photoperiod. Unexpectedly, we found that photoperiod had no role in regulating the life cycle, but diapause was maintained as long as pupae were exposed to cold. Pupae born before February needed a slightly longer exposure to high temperatures to terminate diapause if the cold period was short. Despite the apparent importance of a long period of chilling for life cycle synchrony, it was not required to terminate diapause. This finding of cold mainly preventing, rather than promoting, diapause termination is not novel among temperate insects, but it is rare. Slow diapause termination as a response to exceptionally long exposure to high, not low, temperatures seems to be a cornerstone for synchronizing the life cycle in the deer ked.

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.

Natural variation in Drosophila melanogaster diapause due to the insulin-regulated PI3-kinase

This study links natural variation in a Drosophila melanogaster overwintering strategy, diapause, to the insulin-regulated phosphatidylinositol 3-kinase (PI3-kinase) gene, Dp110. Variation in diapause, a reproductive arrest, was associated with Dp110 by using Dp110 deletions and genomic rescue fragments in transgenic flies. Deletions of Dp110 increased the proportion of individuals in diapause, whereas expression of Dp110 in the nervous system, but not including the visual system, decreased it. The roles of phosphatidylinositol 3-kinase for both diapause in D. melanogaster and dauer formation in Caenorhabditis elegans suggest a conserved role for this kinase in both reproductive and developmental arrests in response to environmental stresses.

Are bottlenecks associated with colonization? Genetic diversity and diapause variation of native and introduced Rhagoletis completa populations

The success of invasive species appears to be a paradox: despite experiencing strong population bottlenecks, invasive species are able to successfully establish in new environments. We studied how the walnut husk fly, Rhagoletis completa, was able to successfully colonize California from the Midwestern United States, by examining genetic diversity and diapause variation of native and introduced fly populations. Climate plays an important role in the successful establishment of introduced insects, because insect diapause is highly dependent upon external climatic conditions. We examined if: (1) fly populations show signs of a population bottleneck, (2) native and introduced flies differ in diapause length when exposed to California and Midwestern climatic conditions, and (3) population genetic diversity is related to variation in diapause length. We assessed if fly diapause conformed more to a model of establishment by local adaptation or to a model of a highly plastic "general-purpose genotype". Our results indicate that only two populations close to the original introduced location showed signs of a population bottleneck, and native and introduced populations did not differ in genetic diversity. Genetic diversity increased in the northern introduced populations, suggesting that multiple introductions have occurred. Flies emerged about 2 weeks earlier under the Midwestern treatment than the California treatment, and introduced flies emerged about a week earlier than native flies. All flies emerged when walnuts are typically available in California. Although variance in diapause length differed between populations, it did not vary between populations or regions. Furthermore, genetic diversity was not associated with diapause variation. Therefore, multiple introductions and a "general-purpose genotype" appear to have facilitated the fly's invasion into California.

Allopatric genetic origins for sympatric host-plant shifts and race formation in Rhagoletis.

Tephritid fruit flies belonging to the Rhagoletis pomonella sibling species complex are controversial because they have been proposed to diverge in sympatry (in the absence of geographic isolation) by shifting and adapting to new host plants. Here, we report evidence suggesting a surprising source of genetic variation contributing to sympatric host shifts for these flies. From DNA sequence data for three nuclear loci and mtDNA, we infer that an ancestral, hawthorn-infesting R. pomonella population became geographically subdivided into Mexican and North American isolates approximately 1.57 million years ago. Episodes of gene flow from Mexico subsequently infused the North American population with inversion polymorphism affecting key diapause traits, forming adaptive clines. Sometime later (perhaps +/-1 million years), diapause variation in the latitudinal clines appears to have aided North American flies in adapting to a variety of plants with differing fruiting times, helping to spawn several new taxa. Thus, important raw genetic material facilitating the adaptive radiation of R. pomonella originated in a different time and place than the proximate ecological host shifts triggering sympatric divergence.

Geographic and parental influences on diapause by a polyphagous insect herbivore

Summary1 Facultative diapause permits insects to initiate an additional generation when conditions are favourable, or to enter diapause when they are not. Although the principle environmental cues for diapause induction are often temperature and photoperiod, we demonstrated recently that the quality of host plant consumed by the larvae ofChoristoneura rosaceana(Harris) (Lepidoptera: Tortricidae) influences the proportion of larvae that enter diapause and therefore voltinism.2 Previous work has also suggested that non‐Mendelian parental effects influence diapause byC. rosaceanalarvae, but the mechanism was not established. Here, we report further evidence for parental effects on diapause and examine whether parental diet influences diapause induction.3 We collected larvae ofC. rosaceanafrom six sites at three latitudes in the Province of Québec. There was an increase in the proportion of larvae entering diapause with increasing latitude. Rearing larvae under identical conditions on artificial diet for two generations significantly reduced variation in diapause among populations within a given latitude, and supported the existence of parental effects that influence offspring diapause.4 Experiments with different host plants and high‐ vs. low‐quality artificial diets provided no evidence that the parental effects on offspring diapause were based on the quality of food consumed by parents. Diapause inC. rosaceanaappears influenced by a hierarchy of cues, including temperature, photoperiod, current larval diet and parental environment.

Oviposition and incubation environmental effects on embryonic diapause in a ground cricket

Maternal effects on offspring phenotype are well known in organisms inhabiting seasonal environments. Mothers that perceive climatological changes indicative of winter's onset will often produce increasing numbers of offspring that enter a state of arrested morphogenesis known as ‘diapause’. In this study of bivoltineAllonemobius sociusground crickets, we manipulated the abiotic environment experienced by ovipositing females (i.e. maternal-oviposition environment) and that experienced by offspring incubating as eggs (i.e. egg-incubation environment) to assess the degree to which mothers contribute to the expression of embryonic diapause in her offspring. Analysis of variance components indicated that variation in maternal-oviposition environment contributed only about 4% to the total variation in diapause incidence, compared to about 24% from embryos responding directly to variation in their incubation environment. Moreover, the 8% contribution from between-family variation was significant, suggesting that parental genes and maternal biotic and abiotic environments contribute to diapause expression in offspring. Although these findings suggest that maternal physi-ology itself contributes little to embryonic diapause variation inA.socius, they do not preclude other maternal behaviours (e.g. placement of eggs at different soil depths) that may affect offspring diapause.

Variation in diapause and sex ratio in the parasitoid Asobara tabida

AbstractAsobara tabida Nees (Hymenoptera: Braconidae) is a widespread parasitoid, attacking larvae of Drosophila (Diptera: Drosophilidae) species in fermenting substrates. In this species, geographic variation is found in the percentage of parasitoids entering diapause and in the sex ratio of emerging parasitoids. Percentage diapause appears to be influenced by host species (more parasitoids enter diapause in D. melanogaster Meigen than in D. subobscura Collin) and temperature. It is not correlated with any of the abiotic factors investigated, but is correlated with survival probability in D. melanogaster larvae and with the time of year in which the experiment was conducted (even though none of the parasitoids experienced natural day light). Sex ratio was only found to correlate with percentage diapause, suggesting that males enter diapause more frequently than females. It is concluded that A. tabida uses diapause to survive both unfavourable abiotic and biotic circumstances.

Altitudinal variation in life cycle syndromes of California populations of the grasshopper, Melanoplus sanguinipes (F.)

Life cycles of California populations of the grasshopper, Melanoplus sanguinipes, varied along an altitudinal gradient. Temperature records indicate a longer season at low altitude on the coast, based on computation of degree days available for development, even though summer air temperatures are cooler than at high altitude; this is a result of warm soil temperatures. At high and low altitudes there was a high proportion of diapause eggs oviposited, while intermediate proportions of diapause eggs occurred at mid altitudes. The low altitude, and especially sea level, populations diapaused at all stages of embryonic development, while at high altitudes most diapause occurred in the late stages just before hatch. Diapause was more intense at high altitudes. One result of diapause differences was delayed hatching in the sea level population. Nymphal development and development of adults to age at first reproduction were both accelerated at high altitude relative to sea level. At lower temperatures (27° C) there was a tendency for short days to accelerate development of sea level nymphs, but not high altitude nymphs. In both sea level and high altitude grasshoppers, short days accelerated maturation of adults to onset of oviposition at warm temperature (33° C); there was little reproduction at 27° C. Population differences for all traits studied appear to be largely genetic with some maternal effects possible. We interpret diapause variation at low and mid altitudes to be responses to environmental uncertainty and variations in development rates to be adaptations to prevailing season lengths.

Larval Diapause Variation in a Tree-Hole Mosquito

Journal Article Larval Diapause Variation in a Tree-Hole Mosquito Get access S. R. Sims S. R. Sims Search for other works by this author on: Oxford Academic Google Scholar BioScience, Volume 32, Issue 5, May 1982, Pages 337–338, https://doi.org/10.2307/1308852 Published: 01 May 1982