The effect of photoperiod on life history and blood‐feeding activity in Aedes albopictus and Aedes aegypti (Diptera: Culicidae)

Several studies have examined how climatic variables such as temperature and precipitation may affect life history traits in mosquitoes that are important to disease transmission. Despite its importance as a seasonal cue in nature, studies investigating the influence of photoperiod on such traits are relatively few. This study aims to investigate how photoperiod alters life history traits, survival, and blood‐feeding activity in Aedes albopictus (Skuse) and Aedes aegypti (Linnaeus). We performed three experiments that tested the effects of day length on female survival, development time, adult size, fecundity, adult life span, and propensity to blood feed in Ae. albopictus and Ae. aegypti. Each experiment had three photoperiod treatments: 1) short‐day (10L:14D), 2) control (12L:12D), and 3) long‐day (14L:10D). Aedes albopictus adult females were consistently larger in size when reared in short‐day conditions. Aedes aegypti adult females from short‐day treatments lived longer and were more likely to take a blood meal compared to other treatments. We discuss how species‐specific responses may reflect alternative strategies evolved to increase survival during unfavorable conditions. We review the potential impacts of these responses on seasonal transmission patterns, such as potentially increasing vectorial capacity of Ae. aegypti during periods of shorter day lengths.     

Temperature and photoperiod effects on dormancy status and life cycle parameters in Aedes albopictus and Aedes aegypti from subtropical Argentina

Aedes albopictus (Diptera: Culicidae) distribution is bounded to a subtropical area in Argentina, while Aedes aegypti (Diptera: Culicidae) covers both temperate and subtropical regions. We assessed thermal and photoperiod conditions on dormancy status, development time and mortality for these species from subtropical Argentina. Short days (8 light : 16 dark) significantly increased larval development time for both species, an effect previously linked to diapause incidence. Aedes albopictus showed higher mortality than Ae. aegypti at 16 °C under long day treatments (16 light : 8 dark), which could indicate a lower tolerance to a sudden temperature decrease during the summer season. Aedes albopictus showed a slightly higher percentage of dormant eggs from females exposed to a short day, relative to previous research in Brazilian populations. Since we employed more hours of darkness, this could suggest a relationship between day‐length and dormancy intensity. Interestingly, local Ae. aegypti presented dormancy similar to Ae. albopictus, in accordance with temperate populations. The minimum dormancy in Ae. albopictus would not be sufficient to extend its bounded distribution. We believe that these findings represent a novel contribution to current knowledge about the ecophysiology of Ae. albopictus and Ae. aegypti, two species with great epidemiological relevance in this subtropical region.     

Photoperiod cues and patterns of genetic variation limit phenological responses to climate change in warm parts of species’ range: Modeling diameter‐growth cessation in coast Douglas‐fir

The phenology of diameter‐growth cessation in trees will likely play a key role in mediating species and ecosystem responses to climate change. A common expectation is that warming will delay cessation, but the environmental and genetic influences on this process are poorly understood. We modeled the effects of temperature, photoperiod, and seed‐source climate on diameter‐growth‐cessation timing in coast Douglas‐fir (an ecologically and economically vital tree) using high‐frequency growth measurements across broad environmental gradients for a range of genotypes from different seed sources. Our model suggests that cool temperatures or short photoperiods can induce cessation in autumn. At cool locations (high latitude and elevation), cessation seems to be induced primarily by low temperatures in early autumn (under relatively long photoperiods), so warming will likely delay cessation and extend the growing season. But at warm locations (low latitude or elevation), cessation seems to be induced primarily by short photoperiods later in autumn, so warming will likely lead to only slight extensions of the growing season, reflecting photoperiod limitations on phenological shifts. Trees from seed sources experiencing frequent frosts in autumn or early winter tended to cease growth earlier in the autumn, potentially as an adaptation to avoid frost. Thus, gene flow into populations in warm locations with little frost will likely have limited potential to delay mean cessation dates because these populations already cease growth relatively late. In addition, data from an abnormal heat wave suggested that very high temperatures during long photoperiods in early summer might also induce cessation. Climate change could make these conditions more common in warm locations, leading to much earlier cessation. Thus, photoperiod cues, patterns of genetic variation, and summer heat waves could limit the capacity of coast Douglas‐fir to extend its growing season in response to climate change in the warm parts of its range.     

Cold adaptation in the Asian tiger mosquito's native range precedes its invasion success in temperate regions

Adaptation to environmental conditions within the native range of exotic species can condition the invasion success of these species outside their range. The striking success of the Asian tiger mosquito, Aedes albopictus, to invade temperate regions has been attributed to the winter survival of diapause eggs in cold environments. In this study, we evaluate genetic polymorphisms (SNPs) and wing morphometric variation among three biogeographical regions of the native range of A. albopictus. Reconstructed demographic histories of populations show an initial expansion in Southeast Asia and suggest that marine regression during late Pleistocene and climate warming after the last glacial period favored expansion of populations in southern and northern regions, respectively. Searching for genomic signatures of selection, we identified significantly differentiated SNPs among which several are located in or within 20 kb distance from candidate genes for cold adaptation. These genes involve cellular and metabolic processes and several of them have been shown to be differentially expressed under diapausing conditions. The three biogeographical regions also differ for wing size and shape, and wing size increases with latitude supporting Bergmann's rule. Adaptive genetic and morphometric variation observed along the climatic gradient of A. albopictus native range suggests that colonization of northern latitudes promoted adaptation to cold environments prior to its worldwide invasion.     

中华通草蛉自然越冬成虫在长、 短光周期下滞育解除中体内相关酶活力变化

光周期信号在昆虫的环境适应中发挥着重要作用, 昆虫能够通过感受光周期的变化来调节体内生理生化过程, 以适应环境的变化。为明确光周期对中华通草蛉Chrysoperla sinica越冬成虫滞育解除过程中酶活力的影响, 本研究测定了长光周期（15L∶9D）和短光周期（9L∶15D）条件下, 成虫体内过氧化氢酶（CAT）、 超氧化物歧化酶（SOD）、 Na⁺K⁺-ATP酶和乳酸脱氢酶（LDH） 4种重要酶活力的变化。结果表明： 中华通草蛉雌、 雄成虫CAT活性在长光周期处理5 d达最高值后呈下降趋势; 短光周期处理CAT活性在处理5 d达最高值, 且高于长光周期处理, 在处理10 d迅速下降至最低值, 且均显著低于长光周期处理的CAT活性（P=0.005）, 后迅速上升并在处理15 d （P<0.05）和20 d （P<0.005）活性显著高于长光周期处理。雌、 雄成虫SOD活性在长光周期下处理10 d达最高值, 且显著高于对照（P<0.001）, 且除处理5 d雄虫SOD活性与对照无显著性差异外（P=0.558）其余处理活性均显著低于对照（P<0.05）。雌成虫长光周期处理5 d 的SOD活性显著低于短光周期（P<0.001）, 其余处理活性均显著高于短光周期; 雄成虫长光周期下处理的SOD活性均高于短光周期下, 且处理5 d （P=0.04）, 15 d （P<0.001）和20 d （P=0.003）的活性差异显著。两种光周期条件下雌、 雄成虫Na⁺K⁺-ATP酶活性随处理时间延长呈上升-下降-上升趋势, 且均显著高于处理0 d成虫酶活性（P<0.001）; 短光周期处理不同时间Na⁺K⁺-ATP酶活性均高于长光周期处理, 且除雄成虫处理15 d无显著性差异（P=0.142）外, 其余均差异显著（P<0.05）。两种光周期条件下雌、 雄成虫LDH活性随处理时间延长呈下降趋势, 且均显著低于对照（P<0.001）。中华通草蛉越冬成虫在长、 短两种光周期条件下体内酶活力的差异可能是影响两种光周期下成虫滞育解除过程中体内不同生化物质含量与生殖状态的重要因子。     

Intermittent character of adult diapause in Aelia acuminata (Heteroptera)

The response to photoperiod of field-collected Aelia acuminata was studied by exposing insects to three photoperiodic regimes. After the completion of diapause the insects do not respond to photoperiod: in mid-February the length of the pre-oviposition period is the same under both long and short photoperiods. After the onset of oviposition, insects again become responsive to photoperiod regardless of the time of year: under long day, oviposition lasts until death, whereas under short day the females cease oviposition after about 3 weeks. After a further 2 or 3 months without ovipositing, these females which are still experiencing short-day conditions resume reproduction for a further 4–7 weeks and long-lived individuals have three oviposition periods. A regulatory mechanism is suggested for the alternation between oviposition and diapause.     

Northward range expansion requires synchronization of both overwintering behaviour and physiology with photoperiod in the invasive Colorado potato beetle (Leptinotarsa decemlineata)

Photoperiodic phenological adaptations are prevalent in many organisms living in seasonal environments. As both photoperiod and growth season length change with latitude, species undergoing latitudinal range expansion often need to synchronize their life cycle with a changing photoperiod and growth season length. Since adaptive synchronization often involves a large number of time-consuming genetic changes, behavioural plasticity might be a faster way to adjust to novel conditions. We compared behavioural and physiological traits in overwintering (diapause) preparation in three latitudinally different European Colorado potato beetle (Leptinotarsa decemlineata) populations reared under two photoperiods. Our aim was to study whether behavioural plasticity could play a role in rapid range expansion into seasonal environments. Our results show that while burrowing into the soil occurred in the southernmost studied population also under a non-diapause-inducing long photoperiod, the storage lipid content of these beetles was very low compared to the northern populations. However, similar behavioural plasticity was not found in the northern populations. Furthermore, the strongest suppression of energy metabolism was seen in pre-diapause beetles from the northernmost population. These results could indicate accelerated diapause preparation and possibly energetic adjustments due to temporal constraints imposed by a shorter, northern, growth season. Our results indicate that behavioural plasticity in burrowing may have facilitated initial range expansion of L. decemlineata in Europe. However, long-term persistence at high latitudes has required synchronization of burrowing behaviour with physiological traits. The results underline that eco-physiological life-history traits of insects, such as diapause, should be included in studies on range expansion.     

Effects of photoperiod on the reproductive condition of Nile grass rats (Arvicanthis niloticus) from an equatorial population

We evaluated the effects of photoperiod on the reproductive condition of male and female Nile grass rats (Arvicanthis niloticus) descended from members of an equatorial population trapped 2°S of the equator. Study animals housed in 12:12 light:dark (LD) cycles were transferred either to short photoperiod (9:15) or long photoperiod (15:9) for 9 weeks (males) or 11 weeks (females), and various reproductive parameters were assessed. We observed no differences between short‐ and long‐day males with respect to plasma concentration of testosterone, testicular mass, seminal vesicle mass, or spermatogenesis. Similarly, we observed no differences between short‐ and long‐day females with respect to oestrous cycles, uterine mass, follicle size, or presence of corpora lutea. Reproductive parameters of male and female A. niloticus housed in short‐ and long photoperiods were similar to those typically observed among animals descended from the same equatorial population and housed in LD 12:12. Thus, photoperiod appears not to elicit changes in reproductive condition among A. niloticus from populations whose native habitat lies within 2° of the equator. These data contrast with the results of other studies indicating that photoperiod alters reproductive condition in A. niloticus populations living >10° from the equator.     

Satyrization and satyrization‐resistance in competitive displacements of invasive mosquito species

Competitive displacements or reductions of resident populations of insects, often effected by a related species, may be caused by a variety of mechanisms. Satyrization is a form of mating interference in which males of one species mate with females of another species, significantly decreasing their fitness and not generating hybrids. Satyrization has been established to be the probable cause of competitive displacements of resident mosquitoes by invasive species, especially of Aedes aegypti by Aedes albopictus, two important vectors of dengue and chikungunya viruses. Mathematical models predict that even low levels of asymmetric mating interference are capable of producing competitive displacements or reductions. Couplings of virgin Ae. aegypti females with Ae. albopictus males effectively sterilize these females through the monogamizing actions of male accessory gland products, but the converse interspecific mating does not impact the future reproduction of Ae. albopictus females. Populations of Ae. aegypti exposed to satyrization quickly evolve resistance to interspecific mating, which is believed to ameliorate reproductive interference from, and promote co‐existence with, Ae. albopictus. The evolution of satyrization resistance among Ae. aegypti in laboratory cages is accompanied by fitness costs, such as reduced fecundity and slower receptivity to conspecific males. Cage experiments and field observations indicate that Ae. albopictus males are capable of satyrizing females of other species of the Stegomyia subgenus, potentially leading to competitive displacements, and possible extinctions, especially of endemic species on islands. Examination of other examples of reproductive interference in insects reveals few parallels to the mechanism and outcomes of satyrization by Ae. albopictus. We conclude by posing the hypothesis that satyrization may favor the ecological success of Ae. albopictus, and suggest many lines for future research on this phenomenon.     

Maintenance and termination of reproductive dormancy in an Australian stink bug, Biprorulus bibax

Reproductive dormancy in female Biprorulus bibax Breddin sampled during March-October in Southern New South Wales was terminated by exposure to 25°C under long (15h) and short (10h) photoperiods. 27% of individuals collected during March-May failed to oviposit under short days. Pre-oviposition period under short photoperiod was greates during March-June (22.4–61.7 d) and shortest during July-October (4.2–14.4 d). Under long photoperiod it was greatest in April and May (20.1–22.3 d) and shortest during June-October (4.9–16.6 d). Longevity ranged from 39.8–89.0 d, with longest lived females being collected in June and held under short days. Fecundity ranged from 51.6–164.4 and was greatest in individuals collected in June-July and held under short days. In females held under long days fecundity was greatest in March and lowest in May. These data indicate that female B. bibax pass the winter in a photoperiodically maintained reproductive diapause.     

Large-Scale Modelling of the Environmentally-Driven Population Dynamics of Temperate Aedes albopictus (Skuse)

The Asian tiger mosquito, Aedes albopictus, is a highly invasive vector species. It is a proven vector of dengue and chikungunya viruses, with the potential to host a further 24 arboviruses. It has recently expanded its geographical range, threatening many countries in the Middle East, Mediterranean, Europe and North America. Here, we investigate the theoretical limitations of its range expansion by developing an environmentally-driven mathematical model of its population dynamics. We focus on the temperate strain of Ae. albopictus and compile a comprehensive literature-based database of physiological parameters. As a novel approach, we link its population dynamics to globally-available environmental datasets by performing inference on all parameters. We adopt a Bayesian approach using experimental data as prior knowledge and the surveillance dataset of Emilia-Romagna, Italy, as evidence. The model accounts for temperature, precipitation, human population density and photoperiod as the main environmental drivers, and, in addition, incorporates the mechanism of diapause and a simple breeding site model. The model demonstrates high predictive skill over the reference region and beyond, confirming most of the current reports of vector presence in Europe. One of the main hypotheses derived from the model is the survival of Ae. albopictus populations through harsh winter conditions. The model, constrained by the environmental datasets, requires that either diapausing eggs or adult vectors have increased cold resistance. The model also suggests that temperature and photoperiod control diapause initiation and termination differentially. We demonstrate that it is possible to account for unobserved properties and constraints, such as differences between laboratory and field conditions, to derive reliable inferences on the environmental dependence of Ae. albopictus populations.     

中华通草蛉自然越冬成虫在两种光周期下滞育解除过程中的生理生化变化

摘要: 光周期在昆虫滞育中起重要作用, 而昆虫光敏感性差异导致了光在昆虫滞育解除过程中作用机制的多样性。为明确光周期在中华通草蛉Chrysoperla sinica (Tjeder)滞育解除过程中的调控机制, 本研究测定了长光周期（15L∶9D）和短光周期（9L∶15D）条件下, 中华通草蛉自然越冬种群成虫在滞育解除过程中的干重、 含水量、 过冷却点（supercooling point, SCP）、 结冰点（freezing point, FP）及蛋白质和糖原含量变化。结果表明： 雌成虫干重在长光照下处理30 d低于短光照下, 其他处理时间的干重均高于短光照下, 但处理间差异不显著（P>0.05）; 雄成虫在长光照下处理5 d的干重高于短光照下, 其余处理时间的干重均低于短光照下, 其中处理30 d 时差异显著（P<0.05）。雌、 雄成虫含水量总体呈上升趋势, 长光照下上升显著（P<0.05）, 短光照下上升不显著（P>0.05）。雌、 雄成虫的SCP、 FP在短光照下处理不同时间均无显著性差异（P>0.05）, 而长光照下表现出先降后升的趋势且处理间差异显著（P<0.05）。成虫蛋白含量在短光照条件下雄虫处理10 d以及雌虫处理15 d均显著低于长光照（P<0.05）, 其他处理之间没有显著性差异。短光照处理成虫体内糖原含量均高于长光照, 且有显著性差异（P<0.05）。多因素方差分析表明, 不同光周期对中华通草蛉成虫滞育解除过程中的SCP、 FP、 糖原变化均有显著性影响（P<0.05）, 而对干重、 含水量及蛋白质含量影响不显著;性别只对成虫FP与糖原有显著影响（P<0.05）; 处理时间对干重、 含水量、 SCP、 FP、 蛋白及糖原含量均有显著影响（P<0.01）。3种因素的交互作用对SCP、 FP、 蛋白及糖原含量均有显著影响, 对干重及含水量影响不显著。初产卵成虫体内物质含量变化测定发现, 长光照（处理10 d）下雌、 雄成虫体内蛋白和糖原含量极显著低于短光照（处理25 d）下（P<0.01）, 雄成虫体内含水量显著高于短光照下（P<0.05）。结果说明, 中华通草蛉自然越冬成虫在长、 短两种光周期下均逐渐解除滞育, 但在滞育解除过程中呈现出不同的生理生化变化。     

Assessment of Aedes albopictus (Skuse) (Diptera: Culicidae) clutch size in wild and laboratory populations

Aedes albopictus (Skuse) is an invasive mosquito species found across the southern U.S. with range expansion into many northern states. Intra‐ and interspecific larval competition have been evaluated for Ae. albopictus with respect to subsequent adult size, immature and adult survivability, and its capacity to vector pathogens as an adult. However, limited data are available on egg production as related to larval rearing conditions. Because Ae. albopictus is a container‐inhabiting mosquito that oviposits in resource‐limited habitats, it is found under variable density‐dependent conditions. Therefore, we examined the impact of specific rearing conditions on Ae. albopictus clutch size and adult body size; comparing the egg production values and wing lengths from known developmental densities to those from field‐collected populations. Field populations varied significantly among collection sites in mean clutch size (23 to 46). These clutch sizes were comparable to the mean clutch sizes of females reared at the larval densities of nine (20 eggs) and three (53 eggs) larvae per 3 ml of water in the laboratory. Field populations experienced density‐dependent effects impacting adult mosquito size. Mosquitoes from the four sample sites had mean wing lengths of 1.99, 2.47, 2.51, and 2.54 mm, which were less than the mean wing length of mosquitoes reared at larval densities of three larvae per 3 ml of water (2.57 mm).     

Differentiation of Photoperiod-Induced ABA and Soluble Sugar Responses of Two Quinoa (Chenopodium quinoa Willd.) Cultivars

Adaptation of quinoa (Chenopodium quinoa Willd.) to new regions demands acclimation to day-length, in addition to a host of other abiotic factors. To further elucidate the effects of photoperiod on development of quinoa, two differently adapted cultivars, Achachino (short day) from Bolivia and Titicaca (day-length neutral), were subjected to continuous long (17.5 h) and short (10 h) photoperiod conditions as well as a shift between the two to trigger possible adaptive mechanisms initiated by changes in leaf soluble sugar and ABA concentration. Our findings show both cultivars responding to an increase in photoperiod with significant increases in soluble sugar concentrations and a simultaneous increase in ABA. However, Titicaca exhibited a much stronger ABA response to increase in photoperiod, whereas the increase for Achachino falls within the range of natural diurnal variation. Achachino also showed increasing sensitivity to long photoperiods throughout all reproductive growth stages, resulting in continued flowering, stem elongation and disruption of seed formation, whereas Titicaca was capable of maintaining full seed set under all the photoperiod conditions. Discernible photoperiod-dependent chlorosis of the lower leaves of Titicaca was observed under long photoperiods compared to short photoperiods, implying multi-faceted adaptive responses to changes in photoperiod which may also involve nitrogen and carbon dynamics. Both ABA and sugar signals are possibly involved in regulating the photoperiod-adaptive capability of each cultivar, leading to pronounced differences in growth and reproductive development patterns between the contrasting cultivars.     

Compensating for climate change–induced cue‐environment mismatches: evidence for contemporary evolution of a photoperiodic reaction norm in Colias butterflies

Anthropogenic climate change alters seasonal conditions without altering photoperiod and can thus create a cue‐environment mismatch for organisms that use photoperiod as a cue for seasonal plasticity. We investigated whether evolution of the photoperiodic reaction norm has compensated for this mismatch in Colias eurytheme. This butterfly’s wing melanization has a thermoregulatory function and changes seasonally. In 1971, Hoffmann quantified how larval photoperiod determines adult wing melanization. We recreated his experiment 47 years later using a contemporary population. Comparing our results to his, we found decreased melanization at short photoperiods but no change in melanization at long photoperiods, which is consistent with the greater increase in spring than summer temperatures recorded for this region. Our study shows that evolution can help correct cue‐environment mismatches but not in the same way under all conditions. Studies of contemporary evolution may miss important changes if they focus on only a limited range of conditions.     

Effects of photoperiod and temperature on reproductive diapause in Ophraella communa (Coleoptera: Chrysomelidae), a potential biocontrol agent against Ambrosia artemisiifolia

Abstract To investigate the seasonal adaptation strategies of Ophraella communa to new habitats, the effects and regulation mechanisms of photoperiod and temperature on the reproductive diapause in a population collected from Changsha, Hunan were examined. Adults showed obvious reproductive diapause, which was regulated by photoperiod and temperature. At 30°C, there was no adult diapause occurring under either long‐day or short‐day conditions; at 25°C the pre‐oviposition period was short and fecundity was high in adult females under L : D 16 : 8 h, whereas under L : D 12 : 12 h, a few females entered reproductive diapause; at 20°C under short‐day conditions, all female adults entered diapause. The pre‐oviposition period was significantly prolonged when the pupae and adults were transferred from long‐days to short‐days, but the day length influence was not obvious when they were transferred only in the adult stage. However, the fecundity dropped greatly no matter whether the photoperiod shifted to short‐days only in the adult stage or whether the shift occurred in both the pupal and adult stage. The fecundity was extremely low when photoperiod shifted from long‐days to short‐days in both pupal and adult stages. This was an indication that the pupal and adult stages were the photoperiod‐sensitive stage for adult reproductive diapause. This was especially true for the photoperiod in the pupal stage, which has a distinctly significant regulative effect on reproductive diapause. Additionally, this article also addresses the reason for different photoperiodic response patterns in reproductive diapause induction between the Changsha strain and the Tsukuba strain (Japan) of O. communa.