The critical field photoperiod inducing ovarian diapause in the cabbage whitefly, Aleyrodes proletella (Homoptera: Aleyrodidae)

ABSTRACT. The ovarian and fat body development of cabbage whitefly, Aleyrodes proletella L., has been determined from field samples taken during the transition from a summer to an overwintering population. Between mid and late September 1983, the state of ovarian development decreases as the proportion of diapausing females increases. This is accompanied by maximum fat body development. Diapausing females emerge from late August onwards, and the critical photoperiod, based upon known larval sensitivity and including civil twilight as part of the photophase, is LD 15.5:8.5h. This is in agreement with laboratory studies. The majority of overwintering females are mated. Very few males survive the autumn.     

甘蓝粉虱入侵中国大陆

甘蓝粉虱是一种危险性入侵害虫,中国大陆于2012年7月在北京市朝阳区首次发现,寄主植物为抱茎苦买菜;翌年8月份又在新疆乌鲁木齐和吐鲁番发现,寄主植物为开花生菜和抱茎苦买菜。文章描述了甘蓝粉虱的危害方式、寄主植物、分布范围、识别特征、天敌种类以及传播扩散方式,并分析了在我国大陆的发展趋势和需要密切关注的区域。     

基于种特异性COI标记的新入侵种甘蓝粉虱快速鉴定技术

【目的】针对粉虱类害虫种类多、体型微小、形态相似、难以准确快速识别的问题,以新入侵我国大陆的甘蓝粉虱 Aleyrodes proletella (L.)为靶标,以田间常见的其他10种/隐种粉虱为参照,采用基于线粒体DNA细胞色素C氧化酶亚基I (mitochondrial DNA cytochrome c oxidase subunit I, mtDNA COI) 基因的种特异性 (species-specific COI, SS-COI) PCR方法,研究其快速分子检测技术。【方法】利用mtDNA COI基因通用型引物LCO-1490/HCO-2198获得甘蓝粉虱及其他常见粉虱的COI序列,根据测序结果设计种特异性SS-COI引物1对（APZYJF/APZYJR）,其扩增片段大小为384 bp,同时对该对引物的种特异性及灵敏性进行检测。【结果】种特异性检验结果显示,该对引物仅对甘蓝粉虱的mtDNA COI基因具有扩增效果,对我国常见的其他种类的粉虱包括温室粉虱 Trialeurodes vaporariorum (Westwood)、柑橘粉虱 Dialeurodes citri (Ashmead)、螺旋粉虱 Aleurodicus disperses (Russell)、双钩巢粉虱 Paraleyrodes pseudonaranjae Martin、非洲伯粉虱 Bemisia afer (Priesner et Hosny)以及烟粉虱B. tabaci (Gennadius)5个隐种（MED, Asia I, Asia II 1, Asia II 3和China 1）等不具有交叉反应扩增能力。灵敏性检验结果显示,该对引物不仅对不同性别的成虫具有良好的扩增效能,对2-4龄若虫甚至单粒卵或单头初孵若虫亦具有同样的扩增能力,其最低检测阈值为14.00±0.37 pg/μL（相当于1/25 600头雌性成虫）。【结论】该技术体系完全可用于甘蓝粉虱的快速准确识别及检测监测,对有效阻截其进一步扩张蔓延意义重大。     

Banker plants promote functional biodiversity and decrease populations of the cabbage whitefly Aleyrodes proletella

In this study, potential banker plant systems against the cabbage whitefly Aleyrodes proletella Linnaeus (Hemiptera: Aleyrodidae) were developed under controlled conditions. The two most promising banker plant systems, that is, the parasitoid Encarsia tricolor Förster (Hymenoptera: Aphelinidae) either with Aleyrodes lonicerae Walker on European columbine (columbine system) or with Trialeurodes vaporariorum Westwood (Hemiptera: Aleyrodidae) on Hokkaido squash (pumpkin system), were further evaluated in the field. Although the pumpkin system produced three times more parasitoids than the columbine system, both banker plants led to an 1.5-fold increase in A. proletella parasitism rates. However, only the pumpkin system increased the abundance of syrphid larvae on cabbage by 61.5% and reduced A. proletella populations on average by 4.4%–25.8% depending on the respective assessment date. In conclusion, the pumpkin system revealed to be a promising (supplementary) control measure against A. proletella. Options for further improvement and standardization of the pumpkin system as well as a potential implementation in cabbage production are discussed.     

Photoperiodism in the cabbage whitefly, Aleyrodes brassicae

ABSTRACT. Reproductive diapause was induced in A. brassicae by short photo-period at 15 and 20°C but was almost prevented at 25°C. The critical photoperiod was LD 153/4: 81/4 at 15°C, giving 60% diapause. Photoperiodic sensitivity was confined to the early immature stages. Food plant quality appeared not to influence the induction or prevention of diapause. Chilling facilitated diapause termination. Overwintering females brought into the laboratory as adults or pupae from the field from September to December took longer before laying their first eggs than females collected in January and February.     

The photoperiodic induction of ovarian diapause in the cabbage whitefly, Aleyrodes proletella (Homoptera: Aleyrodidae)

Female cabbage whitefly, Aleyrodes proletella L., overwinter in ovarian diapause. A scoring system for ovarian development has proved valuable in following the progress of diapause and has revealed a marked difference between the effects of stationary short-day treatment and one involving a transfer from long-day to short-day photoperiods. The former results in a weak inhibition of ovarian development, whilst the latter completely inhibits pre-emergence oögenesis and greatly retards post-emergence development. Therefore, A. proletella may be regarded as a long-day-short-day species. Peak photoperiodic sensitivity is located in the third instar, although some degree of sensitivity is present for most of larval development. The inductive effect of 10 short-day cycles, at the time of peak sensitivity, is not abolished by subsequent long-day treatment.     

Effects of temperature on development of the immature stages of the cabbage whitefly, Aleyrodes proletella (Homoptera: Aleyrodidae)

The duration of development of immature stages of Aleyrodes proletella (L.) decreased with increase in temperature. The rate of the developmental time was smallest in the egg and largest in the second instar. The calculated threshold temperatures of development were approximately equal for the egg (10°) and the fourth-instar larva (10.4°) but differed significantly (P<0.05) from those of the first instar (7.3°), second instar (5°), and third instar (6.5°).

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Zusammenfassung Die Entwicklungszeit des Eistadiums und der Larvenstadiums von Aleyrodes proletella nahm mit zunehmender Temperatur ab. Die relative tägliche Entwicklung war am geringsten beim Ei und am höchsten bei der Zweitlarve. Der berechnete Entwicklungsnullpunkt war ungefähr gleich für das Ei (10°) und für die Viertlarve (10,4°), während er für die Erstlarve (7,3°), für die Zweitlarve (5°) und für die Drittlarve (6,5°) etwas tiefer lag.

     

The regulation of development during diapause in Ephestia elutella (Hübner) by temperature and photoperiod

Diapausing larvae of Ephestia elutella reared at 20°C in short photoperiods (LD 11:13), and then maintained 12 weeks or longer at 5–15°C before transfer to 20 or 25°C, pupated sooner than unchilled controls. At 25°C, all samples kept in long photoperiods (LD 15:9) survived better and pupated faster than similarly treated samples held in short photoperiods (LD 9:15). Samples kept at 20°C after chilling pupated much slower than those at 25°C, and, except after exposure at 5°C, pupated at similar rates at LD 11:13 or 15:9, although mortality was higher at the shorter photoperiod. After exposure at 5°C, larvae required increased day-length as well as increased temperature to hasten pupation whereas after exposure at 10°C most responded to increased temperature only.For samples maintained in slightly heated or unheated outbuildings, the summer emergence was poorly synchronized and males on average emerged ahead of females. Samples moved from the unheated outbuilding to 25°C and long days in the laboratory in early spring, however, pupated quickly and males and females emerged together. A late phase of diapause development thus exists requiring both high temperature and long photoperiods to ensure a prompt resumption of morphogenesis. Spring temperatures in the United Kingdom are seldom high enough to synchronize the completion of diapause.     

Phloem‐specific resistance in Brassica oleracea against the whitefly Aleyrodes proletella

The cabbage whitefly [Aleyrodes proletella L. (Hemiptera: Aleyrodidae)] is becoming a serious pest in Brassica oleracea L. (Brassicaceae) crops. However, almost nothing is known about the interaction of this insect with its host plants. Previous studies have shown differences in the natural occurrence of adults, eggs, and nymphs on the closely related B. oleracea cultivars Christmas Drumhead and Riviera grown in the field. In this study, we aimed to identify the nature of these differences and to gain insight into the resistance mechanisms against A. proletella. We used no‐choice experiments on field‐ and greenhouse‐grown plants to show that the differences between the two cultivars are mainly based on antibiosis (traits that reduce herbivore performance) and not on antixenosis (traits that deter herbivory). This was further supported by laboratory choice experiments that indicated little or no discrimination between the two cultivars based on plant volatiles. We showed that resistance is dependent on plant age, that is, resistance increased during plant development, and is mainly independent of environmental factors. Analysis of probing behaviour revealed that the resistance trait affects A. proletella at the phloem level and that morphological differences between the two cultivars are most likely not involved. We suggest that compounds present in the phloem reduce sap ingestion by the whitefly and that this explains the observed resistance.     

Endocrine changes during diapause development in the cabbage army moth Mamestra brassicae

Many insects undergo diapause to survive adverse seasons. Although the mechanism of diapause induction is the subject of extensive study, that of diapause termination remains poorly understood. In the present study, we show the endocrine processes leading to the termination of pupal diapause in Mamestra brassicae. Diapause of this insect is terminated if the pupae are exposed to a low temperature for several weeks. During this period, the prothoracic glands (PGs) of pupae acquire the potential to secrete sufficient ecdysteroids necessary for inducing adult development. The main endocrine changes observed under the low temperature conditions are: (i) the increase in activity of the PGs in two steps; (ii) the increase in responsiveness of the glands to prothoracicotropic hormone (PTTH); and (iii) two‐step increase in PTTH gene expression in the brain. The timing of the first and second increases in PG activity roughly coincides with that of the two steps of increase in PTTH gene expression, and the timing of the increase in the responsiveness of the PGs to PTTH coincides with the second, larger increase in PTTH gene expression. The ablation of the PGs prior to cooling pupae does not affect the increase in PTTH gene expression, whereas brain removal results in a failure to increase PG activity, strongly suggesting that PTTH is the master regulator of diapause development and termination.     

中国新记录种——忍冬粉虱Aleyrodes lonicerae Walker(半翅目:粉虱科)

在北京房山区上方山的白屈菜Chelidonium majus L.叶背上发现了一种粉虱。这是一种中国新记录种:忍冬粉虱Aleyrodes lonicerae Walker 1852。本文描述了其各期的形态特征及种类鉴别特征,并列出世界已知的11科23种寄主植物,记录了2种寄生蜂:恩蚜小蜂Encarsia sp.和桨角蚜小蜂Eretmocerus sp.nr.emiratus,并提供了粉虱及寄生蜂的生态图。     

Night-interruption experiments and action spectra for dawn and dusk in relation to the photoperiodic clock of the cabbage whitefly,Aleyrodes proletella (Homoptera: Aleyrodidae)

The dark period (scotophase) is the most photoperiodically important part of a light-dark cycle in Aleyrodes proletella. Night-interruption studies have revealed three distinct dark stages: the photosensitive stage 1 lasts for about 3 h after dusk and 1-h light breaks both stop and re-set the photoperiodic clock; stage 2 also lasts about 3 h, but is photorefractory to some degree; stage 3 is photosensitive, but short light breaks do not re-set the clock although a 4-h light break (equivalent to a main photophase) does restore the capacity to respond to a normal critical night length in the post-interruption scotophase.Action spectra revealed peak photoperiodic sensitivity to blue light (410–430 nm) with 50% responses., at 1.5 μWcm⁻² and 2.5 μWcm⁻² for the dusk and dawn peaks respectively. These data are consistent with the view that the photopigment is a carotenoprotein.The results are interpreted in terms of the photoperiodic clock in A. proletella operating on the hour glass principle.     

Photoperiodic responses during larval development and diapause of two geographic ecotypes of the rice stem maggot, Chlorops oryzae

Chlorops oryzae is bivoltine in northern Japan but trivoltine in the southern part of the country. In the bivoltine strain, both the egg and larval stages were found to be sensitive to photoperiod. When the egg stage was exposed to a long-day photoperiod (16L:8D), larval development showed a short-day type response, and mature third-instar larvae entered a summer diapause under a long-day photoperiod (15L:9D). When eggs experienced short days, the first-instar larvae entered a winter diapause under short-day conditions, and the critical photoperiod in the larval stage ranged from about 14L:10D to about 12L:12D as the photoperiod experienced by the eggs increased from 12L:12D to 14L:10D. However, the development of the larvae after overwintering was not influenced by the photoperiod. In the trivoltine strain, larval development was retarded under a 14L:10D photoperiod but not under either shorter or longer photoperiods, when larvae had spent the egg stage under a 16L:8D photoperiod. The critical photoperiod of the larval stage for the induction of a winter diapause in the first instar was about 12L:12D, though it varied to some extent with the photoperiod during the egg stage. Thus, Chlorops oryzae was able to adapt itself to the local climatic conditions by the development of variable and complicated photoperiodic responses.     

Physiological aspects of diapause and cold tolerance during overwintering in Pieris brassicae

Abstract The relationship between diapause-associated metabolic suppression and carbohydrate metabolism linked with cold tolerance was investigated in pupae of Pieris brassicae L. Cold tolerance was assessed by measuring the crystallization temperature (T_c) and by estimates of pre-freeze mortality. Metabolic suppression was measured using ³¹P nmr and carbohydrates by GLC. Sorbitol (a possible cryoprotectant) accumulated from the onset of diapause in October until December reaching a concentration of c. 40 mMolal in both years of the study, but then declined from January until adult eclosion in May. The T_c remained between -23 and -25^oC throughout the winter except for a slight rise before eclosion in May. The absence of a significant T_c suppression is as predicted from the low concentration of sorbitol accumulated. The pre-freeze mortality experiments indicate that pupae are most cold tolerant in the period December-February when sorbitol concentration is high, suggesting an alternative cryoprotective role for sorbitol. Glycogen declined at the beginning of diapause until February after which there was some recovery suggesting that it may be the source of carbon for sorbitol synthesis. Diapause-associated metabolic suppression is evident in the low ³¹P nmr resonances of ATP during November-February compared with non-diapause pupae and diapause pupae soon after pupation. The suppression of metabolism at this time may have a direct role in cryoprotection and by itself (rather than sorbitol) account for the increased pre-freeze cold tolerance. ATP appears to increase slowly from February until a sharp increase occurs shortly before eclosion. Arginine phosphate remains high during diapause until late February-March when it begins a decline which continues until eclosion. A period of change in energy and carbohydrate metabolism is apparent at the same time which may indicate diapause termination and related changes in cold tolerance mechanisms. It is argued that in P.brassicae sorbitol accumulates as a result of metabolic suppression and may have no cryoprotective role. However, for species living in, or colonizing, low temperature environments it is a short evolutionary step to exploit this pathway and accumulate high concentrations of polyols as a specialized cold tolerance strategy.     

On the factors inducing the inhibition of diapause in the progeny of diapause females of Trichogramma telengai

The ability to enter diapause in the progeny of Trichogramma Westw. (Hymenoptera: Trichogrammatidae) females that have undergone diapause is markedly reduced or even completely absent. This maternal inhibition of diapause can result from three types of factors: (i) environmental factors that induce maternal diapause; (ii) maternal diapause itself; and (iii) diapause‐terminating environmental factors. The present study aims to determine the factors that prevent diapause in the progeny of a Trichogramma telengai Sor. generation that has undergone diapause. Different tendencies to enter diapause in the maternal generation are induced by different photoperiods (LD 12 : 12 h, 16 : 8 h and 18 : 6 h) during development of the grandmaternal generation and different temperatures (from 10 to 15 °C) during larval development of the maternal generation. To terminate diapause, prepupae of the maternal generation are kept at 4–5 °C in the dark. To estimate the ability to enter diapause, the progeny generation is incubated at 14 °C. The results suggest that the inhibition of the tendency to enter diapause in the progeny of T. telengai females that have undergone diapause is caused by diapause itself rather than by diapause‐inducing or diapause‐terminating environmental factors. This result can be used to clarify the mechanism of the inhibition of diapause in the progeny of females that have undergone diapause.     

Antennal sensilla of whiteflies: Trialeurodes vaporariorum (Westwood), the glasshouse whitefly,and Aleyrodes proletella (Linnaeus), the cabbage whitefly, (Homoptera : Aleyrodidae). Part 2: Ultrastructure

A transmission electron microscope study of the antennal sensilla of the whitefly Trialeurodes vaporariorum and Aleyrodes proletella (Homoptera : Aleyrodidae) revealed that of the sensilla unique to the antennal flagellum (basiconic, coeloconic and small digitate-tipped sensory pegs), basiconic and coeloconic sensilla occur as subtypes. Four subtypes of basiconic cone sensilla occur on the flagella of T. vaporariorum and 3 on A. proletella. All the subtypes of basiconic sensilla have an ultrastructure typical of olfactory sensilla and probably have a primary olfactory function. Two subtypes of coeloconic sensilla occur on the flagella of both species. Their ultrastructure suggests primarily a chemosensory function. The digitate-tipped sensory peg of both species possesses a triad of neurones which have ultrastructural characteristics similar to the known thermo-/hygroreceptors of other insect species. The other sensilla, which occur on the antennae of the whiteflies, include cheatae, campaniform and subcuticular sensilla, all of which have an ultrastructure typical of mechanoreceptors.     

Effect of photoperiod and temperature on diapause in a Florida strain of the tropical warehouse moth Ephestia cautella

A photoperiodically-controlled diapause of the long-day, short-day type was identified in a brown-winged, yellow-eyed strain of Ephestia cautella (Walker). The proportion of larvae diapausing in very long photoperiods was less than in short photoperiods. The mean critical photoperiod, here defined as that photoperiod giving half the maximum percentage of insects that diapause in response to photoperiod at a given temperature, was between 12 and 13 hr for the long-day reaction at both 20 and 25°C. The principal sensitive phase occurred near the time of the last larval moult. The mean duration of diapause was 2–3 months at 20°C and slightly longer at 25°C. The optimum temperature for diapause development was near 15°C, all larvae pupating within 24 days after a 45-day exposure at this temperature. Diapause could be terminated whenever larvae diapausing at 20°C were exposed to as few as five long (15 hr) photoperiods at 25°C. Long photoperiods at 20°C, or short photoperiods (9 hr) at 25°C were less effective in terminating diapause.     

Antennal sensilla of whiteflies: Trialeurodes vaporariorum (Westwood), the glasshouse whitefly,Aleyrodes proletella (Linnaeus), the cabbage whitefly,and Bemisia tabaci (Gennadius), the tobacco whitefly (Homoptera : Aleyrodidae). Part 1: External morphology

Scanning electron microscopy has revealed the detailed structure of the antennae of three species of whitefly, Trialeurodes vaporariorum, Aleyrodes proletella, and Bemisia tabaci (Homoptera : Aleyrodidae). All 3 species have microtrichia and 5 types of sensilla on the antennae: chaetica, campaniform, basiconic, coeloconic, and pegs with digitate tips, the latter 3 of these being unique to the flagellum. The number and distribution of the sensilla unique to the flagellum vary among the species studied. The cuticle of the basiconic and the coeloconic sensilla is pitted and grooved, respectively, a prerequisite for an olfactory and/or a thermo-, hygroreceptive function. A sexual dimorphism does occur with respect to the position of the basiconic sensilla on flagella segment 5 of A. proletella, but not T. vaporariorum or B. tabaci.     

Photoperiodic and temperature effects on the intensity of larval diapause in Sesamia nonagrioides

Abstract. The intensity of larval diapause in Sesamia nonagrioides Lef (Lepidoptera: Noctuidae) was investigated under laboratory conditions. Newly hatched larvae were exposed to different stationary photoperiods (from LD 7 : 17 h to LD 14 : 10 h), at a constant temperature of 25 °C. Diapause incidence was higher when larvae were exposed to daylengths shorter than the critical value (LD 12 : 12 h), whereas the within‐treatment variation in the larval period appeared to be significantly correlated with the photoperiod applied. The incidences of diapause and the duration of larval development were also measured after exposing larvae to short photoperiods (LD 8 : 16 h, LD 10 : 14 h or LD 12 : 12 h) in combination with various temperatures (20, 22.5 or 25 °C). Although an increase in the incidence of diapause appeared with the lowering of the temperature, no statistical differences were observed in the time needed for pupation within the photoperiodic treatments at the temperatures of 20 and 22.5 °C. Furthermore, when diapausing larvae were transferred to the long photoperiod of LD 16 : 8 h, they immediately proceeded to pupation, regardless of the photoperiod or the temperature to which they had been previously exposed, indicating that there were no differences in the intensity of diapause. Photoperiodic changes from LD 10 : 14 h to LD 12 : 12 h or to LD 14 : 10 h at different larval ages reduced the intensity of diapause with (a) early age of transfer and (b) increase of daylength. By contrast, when larvae were transferred from the long photoperiod of LD 14 : 10 h to shorter, such as LD 10 : 14 h or LD 12 : 12 h, a small increase in the intensity of diapause with the shortening of the daylength was apparent. These results support the hypothesis that insects may compare the duration of the photoperiod and could classify them as either longer or shorter in relation to the critical value.     

Effects of photoperiod and temperature on diapause induction and termination in the swallowtail,Sericinus montelus

Abstract Sericinus montelus overwinters as diapausing pupae. In the present study, the effects of photoperiod and temperature on diapause induction and termination of diapause are investigated. The results obtained demonstrate that high temperature can reverse the effect of short day‐lengths on diapause induction. Under an LD 12 : 12 h photoperiod, all pupae enter diapause at 15, 20 and 25 °C, whereas all pupae develop without diapause at 35 °C. No pupae enter diapause under an LD 14 : 10 h photoperiod when the temperature is above 20 °C. Photoperiodic response curves obtained at 25 and 30 °C indicate that S. montelus is a long‐day species and the critical day‐length is approximately 13 h at 25 °C. At 25 °C, the duration of diapause is shortest when the diapausing pupae are maintained under an LD 16 : 8 h photoperiod and increases under LD 14 : 10 h and LD 12 : 12 h photoperiods. Under an LD 16 : 8 h photoperiod, the duration of diapause is shortest when the diapausing pupae are maintained at 25 °C, followed by 20 and 30 °C, and then at 15 °C. These results suggest that a moderate temperature favours diapause development under a diapause‐averting photoperiod in this species. The duration of diapause induced by an LD 12 : 12 h photoperiod is significantly longer at 25 °C than those at 15, 20 and 30 °C, and is shortest at 15 °C. At 25 °C, the duration of diapause induced by LD 6 : 18, LD 12 : 12 and LD 13 : 11 h photoperiods is similar and longer than 90 days. Thus, the diapause‐inducing conditions may affect diapause intensity and a photoperiod close to the critical day‐length has significant influence on diapause intensity in S. montelus.