Precocious metamorphosis of the aphid Myzus persicae induced by the precocene analogue 6-methoxy-7-ethoxy-2,2-dimethylchromene

For up to three days after being treated with the precocene analogue 6-methoxy-7-ethoxy-2,2-dimethylchromene, adult green peach aphids (Myzus persicae) gave birth to offspring (alate and apterous virginoparae, males) which underwent precocious metamorphosis. Only a few precocious gynoparae and no precocious oviparae were recorded. The precocious aphids became adultoid in the third or fourth-instar as a result of corpus allatum destruction. They were able to develop mature embryos, but could not deposit them because of incomplete differentiation of the reproductive tract. The treatment did not induce the production of alate virginoparae in the experimental clone of Myzus persicae. However, a few males were born late in the reproductive sequence of treated apterae. The corpora allata of the treated adults appeared on histological examination to be unaffected by precocene.     

Inhibition by kinoprene of photoperiod-induced male production by apterous and alate viviparae of the aphid Myzus persicae

Photoperiod-induced male production by apterous and alate viviparae of the green peach aphid. Myzus persicae, was reduced or prevented by treating the aphids with the JH analog kinoprene (Zoecon's insect growth regulator ZR 777). Concomitant with this reduction in the number of males was an increase in the production of females. This was most clearly demonstrated for apterous viviparae that had been raised from birth under weak long-night regimes of 10–10.5 hr dark day, and exposed to 0.1% kinoprene-treated radish seedlings for 2–3 days on reaching the fourth larval instar or adulthood.Male production by alate viviparae, whose apterous mothers were born and raised for 5 days under a long-night regime of 15 hr dark day and then maintained under a short-night regime of 8 hr dark day, was completely inhibited by exposing them to kinoprene-treated plants for 2–3 days when they had reached the fourth instar.It is concluded that the kinoprene treatment affects sex chromosome replication in the oögonia of an aphid so that these germ cells develop parthenogenetically into females (2n = 12 with two X chromosomes) rather than males (2n = 11 with one X chromosome). This may result either from a direct action of absorbed kinoprene on the oögonia or from an indirect action whereby the kinoprene stimulates the insect's hormonal system to produce a hormone that stimulates development of the oögonia into females.     

Male production by apterous viviparae of the aphid Myzus persicae temporarily exposed to different scotoperiods

Male production by apterous viviparae of a holocyclic biotype of the green peach aphid, Myzus persicae, was induced by pre-natal and/or post-natal exposures to a long-night regime of 15 h dark per diem. When the apterae were exposed to three or more long nights immediately after birth and they subsequently developed under a short-night regime of 8 h dark per diem, they produced females (mostly alate viviparae) during the first 8–10 days of reproduction and a high proportion of males thereafter. When the apterae were exposed to two long nights immediately before their birth and to short nights thereafter, they produced relatively few females (mostly alate viviparae), and males were deposited already after 4–6 days of reproduction, i.e. 16–18 days after the first exposure to the long nights. The proportion of males among the progeny of these apterae was highest when the two prenatal exposures comprised scotophases of 11–15 h; under such long-night regimes many aphids switched to producing males exclusively. To achieve this effect, the two long scotophases had to be separated by a photophase of more than 1–2 h. Fewer males were produced and most of the apterae reverted to the production of females (apterous viviparae) when the duration of the two prenatal scotophases was 9 h 45 min-10 h 30 min, or 18 h and longer.One long night of 15–39 h could also induce temporary male production if the aphids were exposed to it late in the 4th larval instar or as teneral adults.     

Embryogenesis and the production of males by apterous viviparae of the green peach aphid Myzus persicae in relation to photoperiod

Apterous viviparae of a holocyclic strain of Myzus persicae were given various numbers of prenatal and postnatal exposures to male-inducing, long-night scotophases (15 hr darkness per diem). The sex of the embryos in various positions within their ovarioles was determined from chromosome preparations and correlated with the sequence with which female and male larvae were deposited. An abnormal follicle was found frequently between the female and male embryos in the ovarioles of aphids raised under long nights.Apterous viviparae exposed to long nights from birth or earlier usually produced several females and then switched over to exclusive male production. The more prenatal exposures to long nights given to the aphids, the fewer females were deposited by them and the earlier they started to produce males. When apterae were continuously exposed to long nights from 2 days or more before their birth, some, and occasionally all, their ovarioles were solely androparous. In general, male-producing apterae deposited considerably fewer larvae (female as well as male) than apterae that produced females exclusively. The 1–2 day pause in larviposition that typically occurs between the deposition of the female and male larvae was attributed to a slower rate of development of the male embryos. The results are discussed in relation to the hormonal changes that may be induced by the photoperiodic treatments.     

Photoperiodic determination of the male and female sexual morphs of Myzus persicae

The chronology of the photoperiodic determination of sexual morphs in holocyclic Myzus persicae was studied in the laboratory by transfer of synchronized batches of aphids between long-day (16 hr) and short-day (10 hr) régimes at a constant temperature of 20°C. The length of exposure to the short-day régime was measured in terms of the number of long dark-periods received by the aphids. The photoperiodic response extended over four generations (P, G₁, G₂, and G₃ respectively). When P generation aphids were given short days from the fourth instar, alate viviparae and males appeared successively in generation G₂, oviparae in G₃. Increasing the number of long dark-periods received during the development of G₂ embryos had a cumulative effect on the number which developed into alate viviparae. Determination of all the first-born G₂ aphids as alatae occurred only if their mothers had been exposed to the short-day régime throughout larval development. Alate viviparae gave birth to oviparae if they received a minimum of 4 long dark-periods, starting from a late stage in their embryonic development. The critical stage for ovipara determination in the G₃ embryo was on the sixth or seventh day after ovulation, more than half-way through embryonic development. G₂ aphids were determined as males before ovulation if their parents received 4 (in some circumstances only 3) long dark-periods. In the clones studied, male determination, once initiated in the G₁ parent, could not be reversed by later back-transfer to the long-day régime.     

Effect of juvenile hormone and β-ecdysone on wing determination in the aphid, Myzus persicae

Juvenile hormone (JH) and analogs have no apparent apterizing effect on Myzus persicae, either prenatally or postnatally. Exogenous JH is deleterious and results in pathological wing deformation, delayed development, and increased mortality. β-Ecdysone has no effect on wing development postnatally. These results are consistent with the view that aphids are normally presumptive apterates with dormant wing rudiments and that wing production means diversion from this basic state.     

Production of males and gynoparae by apterous viviparae of Myzus persicae continuously exposed to different scotoperiods

At 18–19°C, apterous viviparae of a holocyclic strain of Myzus persicae raised from birth under constant scotoperiods of 9 hr 26 min–15 hr darkness per diem deposited apterous viviparae, alate viviparae, and males, in that sequence. Under the shortest scotoperiod (9 hr 26 min) in which males were recorded, only a few were deposited at the end of reproduction. With increasing duration of the scotoperiods, the aphids switched to the exclusive deposition of males progressively earlier in their reproductive lives. Thus more males and fewer females were produced; however the proportion of wingless females increased. Similar trends were recorded when the number of prenatal exposures to each scotoperiod was increased from 0 to 7.Alatae developed into gynoparae at scotoperiods of 10 hr 29 min or longer, and into virginoparae at 9 hr 40 min or shorter scotoperiods. Gynoparae and alate virginoparae, as well as alates that produced both oviparae and viviparae were found at 10 hr and 10 hr 15 min.When apterous viviparae were transferred to scotoperiods of 10 hr 29 min or 15 hr only when they attained adulthood, they also deposited males but only toward the end of their reproductive lives.     

Osmoregulation by the aphid Myzus persicae: A physiological role for honeydew oligosaccharides

The dry matter content of honeydew produced by Myzus persicae feeding on artificial diets increased with increasing sucrose concentrations of the diet. Whereas the diet osmolalities ranged from 828 to 1800 milliosmolal, the honeydew osmolality was relatively constant (about 500 milliosmolal). This osmoregulatory capacity is achieved largely by variations in the mean molecular weight of glucose-containing honeydew oligosaccharides.     

Effects of precocene I and II on flight behaviour in Oncopeltus fasciatus, the migratory milkweed bug

Treatment of newly emerged adult Oncopeltus fasciatus with the corpus allatum inhibitors. 7-methoxy-2,2-dimethyl chromene or 6,7-dimethoxy-2,2-dimethyl chromene (preocene I and II) results in an inhibition of long-term flight (presumed migratory) behaviour in both males and females and inhibition of oögenesis in females. Treatment of reproductive females with precocene briefly stimulates flight behaviour (which ceases in such females as oviposition begins) and then subsequently inhibits it. Oviposition also ceases in such females and oöcyte resorption occurs. Topical application or injections of JH III to precocenetreated animals results in immediate restoration of the tendency to make long tethered flights while corpora cardiaca extract injections, corpora cardiaca implantation, sham implantation, sham injections and sham topical applications were ineffective in restoring prolonged flight behaviour to precocene-treated animals.Restoration of flight by JH III injection was observed within 1 hr after treatment with the hormone. These results indicate that JH is necessary for prolonged flight and presumably migratory behaviour in this species and its stimulatory effect on flight behaviour is immediate. Possible mechanisms of action of the hormone on flight behaviour are discussed.     

Laboratory selection for resistance to sulfoxaflor and fitness costs in the green peach aphid Myzus persicae

Sulfoxaflor is a newly released fourth-generation neonicotinoid insecticide for management of sap-feeding pests that have developed resistance to established insecticide groups. The risk of resistance developing to this pesticide in target pests is unclear. We selected a strain of the green peach aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae), for resistance to sulfoxaflor in the laboratory, which showed 199-fold resistance after 45 generations compared to the starting population. Life table analysis showed that the resistant strain had a fitness of 0.83 compared to the susceptible strain. Adult longevity of the resistant strain was reduced by 9.55% compared to the susceptible strain. The period when adults of the resistant strain produced offspring was reduced by 17.19%, while the mean fecundity of the resistant strain was reduced by 15%. These findings suggest that M. persicae can develop a high level of resistance to sulfoxaflor, but fitness costs may result in a recovery of sensitivity when field populations are no longer exposed to sulfoxaflor.     

The hormonal regulation of the larval diapause in the codling moth, Laspeyresia pomonella (Lep. Tortricidae)

The hormonal control of the facultative diapause of the codling moth has been investigated. The diapause can be divided into 4 phases or periods: (1) diapause induction by short-day conditions (SD) in young larvae, (2) initiation of the diapause in the early last larval instar by a high titre of juvenile hormone, (3) onset and maintenance of diapause with inactivity of the neuroendocrine system, as evidenced by the results of neck-ligation experiments, (4)termination of diapause by the production of ecdysteroid.Diapause-induced larvae pupated after spinning the cocoon, if the state of induction was changed by injection with the anti-juvenile hormone precocene II at the beginning of the last larval instar and subsequent results of neck-ligation experiments, (4) termination of diapause by the production of ecdysteroid. treated with juvenile hormone during the first 1.5 days after the last larval moult and subsequently reared under SD. Under LD, continuous application of juvenile hormone during the last larval instar and after spinning did not prevent the insects from moulting to either a supernumerary larva, a pupa or a larval-pupal intermediate. Termination of diapause, i.e. pupation, was achieved by injecting diapausing larvae with 20-hydroxyecdysone. Although juvenile hormone was found to have a prothoractropic effect in diapausing larvae, no pupal moult could be induced by the application of the hormone. Contrary to the hormonal situation before pupation of nondiapausing larvae, no juvenile hormone could be detected before or during the pupation of larvae after diapause.     

Feeding deterrency of flavonoids and related phenolics towards Schizaphis graminum and Myzus persicae: Aphid feeding deterrents in wheat

A number of naturally occurring flavonoids have been tested for their feeding deterrent activity against two aphid species, Schizaphis graminum and Myzus persicae. Most flavonoids, including a number of dihydrochalcones related to phloretin, showed strong deterrency at concentrations well within the range often found in plants. Flavanone and flavone glycosides showed weak feeding deterrency relative to their corresponding aglycones. S. graminum and M. persicae responded similarly towards the compounds tested. The feeding deterrency of wheat extracts towards S. graminum was confined to the phenolic fraction, which included the flavone tricin. The more polar phenolic fraction showed the strongest feeding deterrency towards S. graminum.     

An aphid circadian rhythm: Factors affecting the release of sex pheromone by oviparae of the greenbug, Schizaphis graminum

Adult oviparae of Schizaphis graminum emit a sex pheromone from scent plaques on the metathoracic tibiae, as do oviparae of other aphid species. By allowing males to walk along a wire walkway, a turning response and an increased rate of antennation were observed when the aphids were within 1–2 cm of a pheromone source. Adult males responded but 4th instar larval males did not.The onset of the release of the pheromone by the oviparae is triggered by the initiation of the light phase in a LD-cycle, and is governed by a circadian rhythm with a free-running period of 25.6 hr under continuous illumination of 15 lx. Daily pheromone release peaks 4–7 hr after lights-on and reaches a maximum on days 6–8 of adulthood.     

Vitellogenin titre in haemolymph of Locusta migratoria in normal adults,after ovariectomy,and in response to methoprene

Vitellogenin in the haemolymph of Locusta migratoria was assayed by rocket immunoelectrophoresis to elucidate aspects of its regulation. In many normal adult females, vitellogenin first appeared on days 5–9, rose quickly to peak levels, and declined before a second vitellogenic cycle; in others, it appeared later and built up more slowly. The timing of first appearance of vitellogenin, and proportions of early and late-developing individuals, differed markedly in groups from the same colony assayed in different years, suggesting effects of both genetic and environmental variation. Average peak levels of vitellogenin were 25–30 mg/ml. After ovariectomy, vitellogenin appeared near the normal time and increased for several weeks to about 300 mg/ml; haemolymph volume also increased greatly, so that the total haemolymph-vitellogenin pool reached about 300 mg/individual, or 100 times the normal amount. After ovariectomy, no cyclicity of vitellogenin accumulation was apparent. These results show that the ovary is not required for stimulation of vitellogenin synthesis, and suggest that normal cycling may depend on inhibition by the mature ovary. Females treated with ethoxyprecocene on day 1 of adult life to inactivate the corpora allata did not produce vitellogenin, but were induced to do so with the juvenile hormone analogue, methoprene. After injection of 150 μg of methoprene in mineral oil, there was one day lag, then vitellogenin increased in the haemolymph to the normal peak level and declined slowly to zero during 5 weeks; after a second injection of methoprene, vitellogenin re-appeared more rapidly, with less lag, reflecting accelerated secondary hormonal stimulation of vitellogenin synthesis in the fat body. Adult males showed no detectable haemolymph vitellogenin even after injection of large doses of methoprene.     

Hormonal control of vitellogenin synthesis in Oncopeltus fasciatus

Previous work indicated the existence of two vitellogenins (A and B) in the haemolymph of Oncopeltus fasciatus, and that vitellogenin B was juvenile hormone (JH)-dependent whereas A was not (Kelly and Telfer, 1977). We have extended these results using several electrophoretic techniques in combination with limited proteolysis of key proteins to show that (1) vitellogenin B is present in eggs in a modified form while vitellogenin A cannot be detected in eggs. (2) Vitellogenin A may be a precursor of B since it has a molecular weight of 200,000D, approximately three times that of vitellogenin B (68,000D) and analysis by limited proteolysis shows that two proteins to be nearly identical. (3) Neither ovariectomy nor treatment with the anti-allatotropin, precocene II prohibits the appearance of vitellogenins A and B in the haemolymph. (4) Injection of ecdysone or 20-hydroxyecdysone into adult, male Oncopeltus fasciatus induces the appearance of both vitellogenin A and B in the haemolymph, suggesting the possible involvement of ecdysteroids in the control of vitellogenin synthesis in this species. (5) We have no evidence for JH control of the synthesis of vitellogenin, however, the ratio of vitellogenin A to B in the haemolymph is higher in the precocene-treated females.     

Parturition and alate morph determination in the aphid Megoura viciae

In reproducing apterae of Megoura viciae, parturition is often completely arrested during periods of isolation from the host plant. In contrast, surgical removal of the rostrum (including the stylets), amputation of the extremities of the legs, or decapitation, all stimulate parturition away from the plant. These operations also induce alata-producing aphids to revert immediately to the production of apterae, but have no detectable effect on aptera-producers. Carbon dioxide or ether anaesthesia and nitrogen narcosis have a similar action on this maternally controlled response. Although the rostrum and tibio-tarsus bear sensilla whose removal might well be involved in inducing parturition, the influence on morph change is probably indirect and is to a great extent associated with the delay in the resumption of parturition. The effect can be reproduced by isolating individual aphids away from the food plant. The morph change cannot, however, be attributed to starvation since it also occurs when the genital pore of an actively feeding aphid is temporarily occluded. The change in physiology appears to be associated with the retention of embryos at a time when there is no sensory input from crowding.

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Résumé Chez les Megoura viciae aptères, la parturition est souvent complètement arrêtée pendant les périodes de séparation de la plante hôte. Par opposition, l'amputation chirurgicale du rostre (y compris les stylets), de l'extrémité des pattes (tarse et une partie du tibia), ou la décapitation, stimulent toutes la parturition en l'absence de la plante. Les pucerons induits à produire des ailés (élevage antérieur en groupe) retournent, après ces opérations, immédiatement vers la production d'aptères. Le dioxyde de carbone ou l'anesthésie à l'éther et la narcose à l'azote ont une action semblable sur ce déterminisme maternel. Bien que l'élimination des sensilla portées par le rostre et par l'ensemble tarse-tibia puisse être déterminante dans l'induction de la parturition, l'action sur le changement de type semble être principalement associée au retard consécutif dans la reprise de la parturition. Les anesthésiants qui, eux aussi, retardent l'apparition ou la reprise de la parturition, ont probablement une action indirecte du même type.Les pucerons groupés, isolés de la plante hôte pendant plus de 24 h, ont aussi tendance à retourner immédiatement à la production d'aptères. Ce changement de type ne peut, cependant, être attribué au jeûne puisqu'il se produit aussi quand, chez un puceron s'alimentant activement, le pore génital est momentanément bouché. Le changement physiologique semble être associé à la rétention des embryons à un moment où il n'y a pas l'influence sensorielle du groupement.Aucun de ces traitements, à l'exception du groupement, n'induit des pucerons antérieurement isolés à devenir des producteurs d'ailés.

     

Juvenile hormone esterases in diapause and nondiapause larvae of the European corn borer, Ostrinia nubilalis

Haemolymph levels of juvenile hormone esterase, 1-naphthyl acetate esterase, and juvenile hormone were measured in synchronously staged diapause and nondiapause larvae of the European corn borer, Ostrinia nubilalis. Juvenile hormone esterase levels were monitored using juvenile hormone I as a substrate while juvenile hormone titres were measured with the Galleria bioassay. Haemolymph of nondiapause larvae showed two peaks of juvenile hormone hydrolytic activity: one near the end of the feeding phase and a smaller one just prior to pupal ecdysis. These peaks of enzyme activity correlated well with the low levels of haemolymph juvenile hormone. Juvenile hormone titres were high early in the stadium then showed a second peak during the prepupal stage coinciding with low esterase activity. Diapause haemolymph had peak juvenile hormone esterase activity nearly 4 times the nondiapause level, reaching a peak near the end of the feeding phase. Diapause-destined larvae retained high juvenile hormone titres even during the rise of the high esterase levels. 1-naphthyl acetate esterase levels did not correlate with the juvenile hormone esterase levels in either the diapause or nondiapause haemolymph. High levels of 1-naphthyl acetate esterase activity were associated with moulting periods.     

Action spectra for the photoperiodic control of polymorphism in the aphid Megoura viciae

Energy compensated action spectra are given for the photoperiodic control of polymorphism in Megoura. The production of ‘long day’ parthenogenetic virginoparae and ‘short day’ oviparae mainly depend on the night length. Light has three different effects. ‘Early’ interruptions of the dark phase in a long night cycle reverse the time-measuring dark response. The action spectrum for a 1 hr interruption placed 1.5 hr after the onset of darkness (during dark stage 1) shows a relatively narrow band of activity, mainly in the blue (450–470 nm). The threshold is ca. 0.25 μW cm^?2. ‘Late’ interruptions placed 7.5 or 8 hr after dark hour 0 (during dark stage 3) strongly promote the production of virginoparae without causing a reversal of the response. The action spectrum has the same blue maximum but sensitivity extends into the yellow and red spectral regions. The third photosensitive component, the main photoperiod itself, is required for initiating the dark timing response and has an intermediate action spectrum. Time/intensity curves for a single wavelength (471 nm) show that the responses during stages 1 and 3 depart markedly from reciprocity. Short durations cannot be compensated by high intensities. The shape of the reciprocity curve for an ‘early’ interruption suggests that the stage 1 response is complete after ca. 1.25 hr.The action spectra are believed to be compatible with the view that the photoreceptor is a caroteno-protein. It is suggested that all three pigment forms are related and that time measurement is largely a function of spontaneous ‘dark reaction’ changes in the pigment system. Stage 1 may represent the reversible conjugation phase of the protein/chromophore moieties. In Stage 2 the pigment is presumably photorefractory and is transitional to the highly sensitive broad spectrum form of stage 3.     

The source and action of head factors regulating juvenile hormone esterase in larvae of the cabbage looper, Trichoplusia ni

Brain (median or lateral regions) or suboesophageal ganglion (SOG) homogenates of Day 1 fifth instar larvae of Trichoplusia ni induced the appearance of haemolymph juvenile hormone esterase (JHE) when injected into Day 1, Day 2 or early Day 4 fifth instar ligated hosts. Brain and SOG homogenates of late fourth instars also induced JHE when injected into Day 1 hosts, whole late fifth instar and pupal tissue did not. The pattern of JHE induction by early fourth through Day 3 fifth instar brain and SOG homogenates correlated with natural haemolymph JHE activity occurring at these times. Implantation of late fourth and Day 1 fifth instar brains and/or SOG into similar age hosts similarly induced JHE activity while prothoracic and abdominal ganglia did not. The relative levels of induction following implantation were SOG<brain<brain+SOG. JHE activity which appears in the haemolymph following injection of brain homogenates appears to be largely due to a single enzyme which has an isoelectric point indistinguishable from that of the natural haemolymph enzyme. Evidence is presented which suggests that inhibitory as well as stimulatory brain factors are involved in JHE regulation.     

Regulation of juvenile hormone esterase activity in the European corn borer, Ostrinia nubilalis

The regulation of juvenile hormone esterase in last-instar diapause and nondiapause larvae of Ostrinia nubilalis was investigated using topically applied juvenile hormone I and a juvenile hormone mimic, methoprene. The influence of the head on juvenile hormone esterase was also investigated. Both juvenile hormone and methoprene caused increases in esterase levels when applied to feeding animals. Neither the hormone nor methoprene was capable of elevating nondiapause esterase activity to levels comparable to those found in untreated prediapause larvae. The esterase levels could be elevated in the larval body, without the head, during prepupal development of nondiapause larvae and in post-feeding diapause larvae. In both cases, juvenile hormone or methoprene induced juvenile hormone esterase activity in head-ligated animals. Topically applied methoprene prolonged feeding and delayed the onset of diapause. When methoprene was applied to larvae that had entered diapause, it disrupted diapause by inducing a moult.