Induction of mictic females in the rotifer Brachionus: oocytes of amictic females respond individually to population-density signal only during oogenesis shortly before oviposition

1. One at a time during the reproductive period of amictic females, oocytes fill with yolk and undergo a mitotic maturation division (oogenesis), are oviposited as single cells, and then develop parthenogenetically into females. Sexual reproduction in Brachionus and several other genera is initiated when amictic females are crowded and oviposit some eggs induced to differentiate into mictic females. Mictic females produce haploid eggs that can develop parthenogentically into males or be fertilised and develop into diapausing embryos called resting eggs. 2. This study examines the time when oocytes in amictic females respond to maternal population density. Is the fate of all oocytes in the germarium irreversibly determined during the early postnatal life of the mother, or is each oocyte labile until just before oviposition? In the former case, the probability of an amictic female producing a mictic daughter at any time throughout her reproductive period would reflect the population density she experienced while young and not that at the time she oviposited an egg. 3. Amictic females of two clones of a Florida strain of B. calyciflorus were cultured singly from birth at a low or high density (in a large or small volume) until about halfway through their reproductive period and then switched (experimental treatment), or not (control treatment), to the other density condition. The results indicate that the female fate of an oocyte is determined by maternal population density during oogenesis. Eggs oviposited soon after transfer from low to high density had the same, or a higher, probability of becoming mictic females compared with those produced by control females kept at the high density; eggs oviposited after transfer from the high to the low density had the same low probability of becoming mictic females as those produced by control females kept at the low density. 4. Control females kept at the high density were less likely to produce mictic daughters as they aged. This decline is not because of a decreased propensity of older females to respond to crowding, as older females responded maximally when transferred from a low to a high population density. 5. As oocytes in amictic females respond to maternal population density only during oogenesis, there is a negligible lag between the population‐density signal in the environment and the commitment to sexual reproduction. This minimises the obligatory two‐generation lag between this signal and production of resting eggs, and thus reduces the possibility that crowding will lead to food limitation before production of these eggs.     

Diapause and overwintering of the hoverfly Episyrphus balteatus

The overwintering biology of the hoverfly Episyrphus balteatus (DeGeer) (Diptera: Syrphidae) was studied and analysed. In Europe, females of this species can overwinter as adults in a facultative, reproductive diapause. The diapause phase is characterized by the ovaries ceasing to develop and by hypertrophy of the fat body. Diapause was induced during the second and third larval instars. The critical photoperiod for inducing diapause was between 11.8 and 11.9 h, corresponding to the day lengths that occur during mid‐ to late September in Hannover, Germany. When temperatures were lower, insects could be induced into diapause at longer day lengths, similar to those that occur in early September in Hannover. A semifield study was done during the winter of 1999/2000 to confirm the results obtained under laboratory conditions and to obtain additional information on the over‐wintering development and mortality of E. balteatus. The results suggested that mortality was correlated with the duration of the experiment and with humidity, rainfall, and temperature. The importance of these results for the regulation of cereal aphid populations by this hoverfly is discussed.     

Cold hardiness in the black rice bug, Scotinophara lurida

Abstract.  The mechanisms and strategies for winter survival of the black rice bug Scotinophara lurida are investigated along with the relationship between cold hardiness and diapause. The ability of S. lurida to survive subzero temperatures varies depending on developmental stage, temperature and exposure duration. Mean supercooling point (SCP) varies from –7.6 to –10.7 °C with developmental stage, but is not significantly different between stages examined. The SCP also varies with season, being lowest in January and increasing rapidly in February and remaining almost at the same level (–7.3 to 9.6 °C) until April The osmolality of haemolymph of field-collected S. lurida adults rises dramatically from 53.9 mOsm kg⁻¹ in August to 75.3 mOsm kg⁻¹ in December, and then declines linearly to 57.0 mOsm kg⁻¹ in May. Field-collected S. lurida adults show a peak glucose content in October, glycerol content in November and trehalose content in December. Only trehalose content decreases after the application of the juvenile hormone analogue, fenoxycarb, suggesting that trehalose is a cryoprotectant during diapause. These various physiological and biochemical traits related to cold tolerance in S. lurida may be, at least in part, under the control of juvenile hormone through the reproductive diapause programme.     

Alternative life cycles controlled by temperature and photoperiod in the oligophagous bug, Dybowskyia reticulata

Abstract. Effects of temperature and photoperiod on the induction and re-induction of adult diapause were examined in Dybowskyia reticulata (Dallas) (Heteroptera: Pentatomidae). Adults collected from the field after overwintering in early summer continued oviposition under long-day conditions of LD 16:8 h at 20 or 25°C, while they re-entered diapause under short-day conditions of LD 12:12 h at 25, 27.5 or 30°C. By contrast, adults reared in the laboratory from eggs at 20 or 25°C entered diapause under both long-day and short-day conditions, whereas those reared at 27.5 and 30°C entered diapause only under short-day conditions. Under quasi-natural conditions in 1993, when summer temperature was low, most adults of the first generation entered diapause in late July. However, in the warmer summer of 1996, oviposition was recorded in many females that ecdysed into adults from July to early August. Even though the seeds of the host plants occur in a restricted period from early summer to early autumn, in warmer years D. reticulata may produce a second generation. The response to temperature with a threshold between 25 and 27.5°C in D. reticulata brings about a switch between the univoltine and bivoltine life cycles.     

二化螟滞育生物钟蛋白TIME-EA4基因的克隆及时空和温度诱导表达分析

【目的】生物测时蛋白TIME-EA4是昆虫特有的一种滞育生物钟蛋白。本文旨在克隆二化螟Chilo suppressalis TIME-EA4基因,研究其在二化螟不同发育时期和不同组织中的时空表达模式及在不同田间种群和不同温度诱导下的滞育和非滞育种群中的表达水平。【方法】通过RACE技术克隆二化螟TIME-EA4基因全长c DNA序列,利用荧光定量PCR的方法检测TIME-EA4基因在二化螟不同发育时期、6龄幼虫不同组织中的表达模式,及在不同季节采集的田间种群和不同温度处理1h后的滞育和非滞育种群6龄幼虫中的表达量变化。【结果】从二化螟中克隆获得TIME-EA4的c DNA序列(Gen Bank登录号:KU356855),全长821 bp,开放阅读框516 bp,编码172个氨基酸;TIME-EA4基因在二化螟不同发育时期和不同组织中均有表达,且在早期蛹、雌雄成虫以及6龄幼虫的头部、脂肪体和中肠中表达水平较高,同时在滞育幼虫组织中的表达高于非滞育幼虫。此外,TIME-EA4基因在滞育种群中的表达量是非滞育种群的3倍左右,同时在非滞育种群中,该基因的表达明显受10℃及以下低温的诱导,且在10℃时表达量最高,而在滞育种群中该基因的表达受温度影响不明显。【结论】结果说明TIME-EA4基因与二化螟的抗低温胁迫以及低温滞育相关。本研究为TIME-EA4在农业害虫滞育的分子作用机制研究奠定了基础。     

Persistence of a Wolbachia infection frequency cline in Drosophila melanogaster and the possible role of reproductive dormancy

Field populations of arthropods are often polymorphic for Wolbachia but the factors maintaining intermediate Wolbachia frequencies are generally not understood. In Drosophila melanogaster, Wolbachia frequencies are highly variable across the globe. We document the persistence of a Wolbachia infection frequency cline in D. melanogaster populations from eastern Australia across at least 20 years, with frequencies generally high in the tropics but lower in cool temperate regions. The results are interpreted using a model of frequency dynamics incorporating cytoplasmic incompatibility (CI), imperfect maternal transmission and Wolbachia effects on fitness. Clinal variation is less pronounced in eastern North America which may reflect annual recolonization at higher latitudes. Limited samples from Africa from latitudes matching our tropical and subtropical samples from Australia and North America show comparably high infection frequencies, but some equatorial samples show lower frequencies. Adult dormancy across cold periods may contribute to the Australian Wolbachia cline. Infected flies exposed to cold conditions for an extended period had reduced fecundity and viability, an effect not evident in unexposed controls. These fitness costs may contribute to the relatively low Wolbachia frequencies in Australian temperate areas; whereas different processes, including CI induced by young males, may contribute to higher frequencies in tropical locations.     

Moisture is required for the termination of egg diapause in the chrysomelid beetle, Homichloda barkeri

Four factors (moisture, light regime, temperature, food type) were examined for their effects on the embryonic diapause of Homichloda (Weiseana) barkeri (Jacoby) (Coleoptera: Chrysomelidae), a biocontrol agent for prickly acacia, Acacia nilotica (L.) Willdenew ex Delile (Mimosaceae). Moisture is critical for termination of diapause. A single wetting of eggs resulted in a low hatch rate while a sequence of wetting events followed by periods of dryness produced a high hatch rate. A relatively constant proportion of embryos within each batch initiated development at each wetting event, with hatching complete after the eighth wetting event in these trials. An extended interval between wetting events, tested at up to 23 days, did not result in a decreased overall hatch rate. A threshold time of exposure to moisture of between 3 to 6 h is required before development proceeds. The response of eggs to the moisture regime is seen as a strategy for taking advantage of available food after rainfall by terminating diapause, rather than merely a quiescent response to the absence of moisture. Temperature affected development time and the proportion of eggs that developed. Experimental manipulations of photoperiod and host-plant availability showed no effect on embryonic development.     

Selection for non-diapause in the bumblebee Bombus terrestris, with notes on the effect of inbreeding

Bumblebees are generally believed to be annual insects. However, here we will show that under laboratory conditions the bumblebee Bombus terrestris (L.) can produce a second generation without a period of cold storage (diapause) or CO₂ narcosis (a method to break diapause). It is also shown that this so-called non-diapause trait can be selected for. The percentage of non-diapausing queens increased from 8% (minimum) to 97% (maximum) in two generations of selection. However, it was not possible to maintain isofemale non-diapause lines. Colonies of the fourth and fifth generation remained small (expressed in worker number) and produced a small number of queens. Also the percentage of queens that started laying eggs (defined as the percentage non-diapause) decreased in the fourth and fifth generations. To study whether this decline of the non-diapause lines was caused by inbreeding, a control experiment was conducted. In this control experiment queens were mated with their brothers (full-sib mating) for several generations and the number of queens that start egg laying was measured. This revealed that inbreeding can have a negative effect on the egg-laying capacities of queens thus causing the decline of inbred (non-diapause) lines.     

Predicting spring emergence of blackcurrant leaf midge (Dasineura tetensi) from air temperatures

The mean duration of post-diapause development of overwintered Dasineura tetensi larvae (in cocoons) was 72.8 (SD=11.4), 45.9 (SD=8.6), 28.7 (SD=6.0), 15.9 (SD=4.3), 10.4 (SD=1.9) and 10.2 (SD=1.8) days at constant temperatures of respectively 10, 12.5, 15, 17.5, 20 and 25 °C in the laboratory. No perceptible development occurred at 5 or 7.5 °C and complete mortality occurred when larvae were held at 30 °C for prolonged periods. The relationship between development rate (r days^–1) and temperature (T °C) was sigmoidal between 10 and 25 °C, the logistic equation r=0.0158+0.085/(1+exp(–0.696(T–17.0))) accounting for 98% of the variation. Larvae entered the winter in diapause. Populations of cocoons were greatest in the surface soil in the centre of bushes adjacent to the crown, 69, 15, 9 and 6% of cocoons occurring in the top 0–1, 1–2, 2–3 and 3–4 cm of the soil, respectively. The time of termination of diapause in the field varied greatly between individuals and from season to season but a significant proportion (>40%) had broken diapause by the end of January in each of the three seasons studied. Diapause was not terminated in the laboratory by chilling over-wintered larvae in cocoons at –2.5, 2.5 or 10 °C for up to 28 days nor when held in a L16:D8 photoperiod. A computer-based phenological forecasting model was constructed using the development rate values (using the INSIM software developed at The Agricultural University, Wageningen, The Netherlands). The model accumulated daily development amounts calculated from daily maximum and minimum air temperatures from 1 February, the end of the coldest period of the year on average and before significant post-diapause development occurred. The model uses boxcar trains to simulate dispersion. The model predicted the time of first emergence of D. tetensi adults in spring at HRI-East Malling generally to within 6 days of the observed time of emergence, and to within 11 days at worst. There was poorer agreement between observed and predicted times of emergence when daily maximum and minimum soil temperatures (depth ca. 3 cm) were used. The use of the model to time insecticidal sprays in relation to the flowering time of blackcurrant is discussed.     

Interpopulation variation of hibernal-aestival-diapause in the egg parasitoid wasp Anagrus takeyanus: adaptation to seasonal host-plant alternation of the tingid host, Stephanitis takeyai

The egg parasitoid wasp, Anagrus takeyanus Gordh & Dunbar (Hymenoptera: Mymaridae), is one of the few natural enemies of the andromeda lace bug, Stephanitis takeyai Drake & Maa (Heteroptera: Tingidae). Studying this wasp in two habitats, I found that at Kyoto, where the bug alternates its host plants seasonally, most wasp individuals in the overwintering eggs of the bug entered summer diapause immediately after winter diapause. However, at Nara, where the bug does not alternate its host plant, the wasp proved to have no summer diapause. Changes in the parasitization rate between generations differed at the two study sites. In Kyoto, the overwintered generation suffered the highest parasitism among the three generations of the year. In contrast, the first and second generations suffered higher parasitization rates in Nara. This difference in level of parasitization between the two study sites corresponded to the different diapause regime of the wasp and the seasonal population trends of the lace bug. Laboratory experiments showed that short photoperiod in combination with low temperature could terminate the long diapause at the time the non-aestivating individuals terminate winter diapause. As a life cycle without host plant alternation is likely to be an ancestral character of the bug, the aestivation of the wasp is thought to have been appended to overwintering as an adaptation to the evolution of seasonal host plant alternation in the lace bug.