Interspecific competition between the polyembryonic wasp Copidosoma floridanum and the gregarious endoparasitoid Glyptapanteles pallipes

We examined interspecific competition between the egg-larval polyembryonic parasitoid Copidosoma floridanum (Hymenoptera: Encyrtidae) and the gregarious larval endoparasitoid Glyptapanteles pallipes (Hymenoptera: Braconidae). Approximately 72% of multiparasitized Acanthoplusia agnata (Lepidoptera: Noctuidae) produced C. floridanum adults, 14% produced G. pallipes adults, and 14% died without producing any parasitoids. Development was delayed and the rate of weight gain was reduced in multiparasitized hosts that produced C. floridanum compared with singly-parasitized hosts. In contrast, only weight was reduced in multiparasitized hosts that produced G. pallipes adults. Compared with single parasitism, the brood size of wasps emerging from multiparasitized hosts was reduced in both species. The percentage of hosts containing precocious larvae of C. floridanum, which are considered to be soldiers, did not increase in response to parasitization by G. pallipes. However, developmental cessation and death of G. pallipes eggs and larvae may be closely related to the number of coexisting precocious larvae.     

Development of the polyembryonic parasitoid Copidosoma floridanum in Trichoplusia ni

Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae) parasitized by the polyembryonic egg-larval parasitoid Copidosoma floridanum (Ashmead) (Hymenoptera: Encyrtidae) attained significantly larger final weights and head capsule widths than unparasitized controls. The difference in weight between parasitized and unparasitized hosts was not entirely accounted for by the weight of the C. floridanum brood. The head capsule widths of all parasitized and unparasitized fifth instars used in the study exceeded the critical threshold of 1.66 mm previously established for T. ni metamorphosis. The critical ratios associated with each T. ni instar of: 1) maximum weight within the instar:head capsule width and 2) maximum weight within the instar:weight at the beginning of the instar differed between parasitized and unparasitized larvae. Development of C. floridanum was synchronized with that of its host. Germ band formation and gastrulation of morulae destined to produce reproductive larvae invariably coincided with the host molt to the ultimate, fifth instar. Reproductive larvae had two instars. Eclosion from the egg to the first instar occurred during day 2 of the host's fifth instar, and ecdysis from the first to the second instar was synchronized with host cocoon spinning. Conversely, embryogenesis of morulae destined to produce precocious larvae began during the host first instar, continued through the second and third instar and ceased during the penultimate, fourth instar. Precocious larvae never molted and died when the host was consumed by the reproductive larvae.

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Résumé T. ni Hübner parasité par le parasitoïde ovo-larvaire C. floridanum Ashmead à développement polyembryonnaire atteint un poids final signficativement plus élevé avec une capsule céphalique plus grosse que les témoins non parasités, sans subir de mues surnuméraires. La différence de poids entre noctuelles parasitées ou non ne correspondait pas entièrement au poids des C. floridanum. Les largeurs des capsules céphaliques de tous les T. ni du cinquième stade dépassaient toutes le seuil critique de 1,66 mm lié à la métamorphose, mais les seuils critiques de taille du corps:largeur de la capsule céphalique et/ou taille et corps, taille initiale du corps au début du stade associé à la mue, différaient chez T. ni parasités ou non. Les développements de T. ni et de C. floridanum étaient synchrones. La formation de la bande germinative et la gastrulation de la morula produisant la multiplication des larves ont coïncidé invariablement avec la mue de l'hôte donnant le dernier stade. Les larves polyembryonnaires ont présenté deux stades. L'éclosion des oeufs s'est produite le deuxième jour du cinquième stade de T. ni, et le passage du premier au second était synchrone de la formation du cocon de l'hôte. Réciproquement, l'embryogenèse de la morula qui donnait des larves précoces commençait pendant le premier stade de l'hôte et se poursuivait à travers les second et troisième, pour cesser pendant le quatrième et pénultième stade.

     

Ultrastructural changes in corpora allata during a cycle of juvenile hormone biosynthesis in embryos of the viviparous cockroach, Diploptera punctata

We have observed changes with time in the fine structure of corpora allata (CA) during a known cycle of increasing and decreasing juvenile hormone (JH) synthesis in late embryos of Diploptera punctata. A previous report showed that rates of JH release were relatively low in 28-day-old embryos, but CA activity subsequently rose linearly to a peak on about day 42, and thereafter steadily declined to a low level on day 64 just before birth (Holbrook et al., 1997). We now show that, regardless of rate of JH synthesis, CA cells are large and replete with organelles which nevertheless exhibit variable morphology in embryos of different age. Highly active CA cells on day 40 contain abundant ring-form mitochondria, whereas CA cells of low activity on days 28 and 64 contain mitochondria that are rod-shaped or globular. Mitochondrial cristae were scarce and indistinct on day 28 but numerous and well developed on day 64. Endoplasmic reticula (ER) are rare on day 28 and appear in increasing numbers when CA activity rises. On day 40, ER are abundant and often exhibit a whorl-like appearance which is not observed on day 28. After day 44, when biosynthetic activity is declining, whorls of ER gradually decrease in number and are ultimately replaced by vesicular smooth ER on day 64. Neurosecretions are found only after day 38, by which time rates of JH synthesis have increased substantially from those of day 28. Except for membranous autophagic vacuoles, which are frequently found when ER whorls disintegrate as rates of JH synthesis decline toward birth, most autophagic vesicles such as multivesicular vesicles and dense bodies occur only sporadically among CA cells at all examined ages. We conclude that synchronous autophagy of exhausted organelles, which results in atrophy of CA cells and long-term arrest of JH synthesis in adult females of D. punctata, does not occur in embryos. The slow cyclic change in rate of JH synthesis in embryonic CA is most likely due to asynchronous autophagic activity and to alterations in certain unique features of intracellular organelles.     

Sex differences in the protection of host immune systems by a polyembryonic parasitoid

Endoparasitoids have the ability to evade the cellular immune responses of a host and to create an environment suitable for survival of their progeny within a host. Generally, the host immune system is suppressed by endoparasitoids. However, polyembryonic endoparasitoids appear to invade their hosts using molecular mimicry rather than immune system suppression. It is not known how the host immune system is modified by polyembryonic endoparasitoids. Using haemocyte counts and measurement of cellular immune responses, we evaluated modification of the host immune system after separate infestations by a polyembryonic parasitoid (Copidosoma floridanum) and another parasitoid (Glyptapanteles pallipes) and by both together (multi-parasitism). We found that the polyembryonic parasitoid maintains and enhances the host immune system, whereas the other parasitoid strongly suppresses the immune system. Multi-parasitization analysis revealed that C. floridanum cancelled the immune suppression by G. pallipes and strengthened the host immunity. This enhancement was much stronger with male than with female C. floridanum.     

The effect of juvenile hormone on temporal polyethism in the paper wasp Polistes dominulus

Juvenile hormone (JH) has an important role in the behavior of eusocial Hymenoptera. Previous work has shown that JH influences aggression and dominance behavior in primitive eusocial insects that lack discrete queen and worker castes (e.g. Bombus bees and Polistes wasps). In contrast, JH is one of the factors that mediates temporal polyethism among workers in advanced eusocial insects that have reproductive castes (e.g. Apis bees and Polybiawasps). Therefore, initial observations suggest that JH may have different roles in primitive and advanced eusocial taxa. Here, we use detailed behavioral observations of marked individuals to test whether JH influences temporal polyethism in the primitive eusocial wasp Polistes dominulus. First, we show that workers in P. dominulus have an age-related division of labor, as workers switch from nest work to foraging as they mature. Then, we show that application of JH accelerates the onset of foraging behavior.Workers treated with JH start foraging at a younger age than control workers. Therefore, JH mediates temporal polyethism in the primitively eusocial insect Polistes dominulus. Received 23 April 2008; revised 6 August 2008; accepted 11 August 2008     

The role of juvenile hormone esterase in terminating larval feeding and initiating metamorphic development in Trichoplusia ni

This study shows, first, that when JH degradation by JHE is blocked with an inhibitor (EPPAT, O-ethyl-S-phenyl-phosphoramidothiolate), prothoracicotropic/ecdysone release/effects are postponed in the cabbage looper Trichoplusia ni (Hübner) (Noctuidae). Thus, JHE is an important component of JH degradation, implying that without normal degradation the JH titer will become abnormally high. Second, this accumulation of endogenous JH in EPPAT treated larvae results in an extra larval molt. Therefore, JHE is also important in the control of the nature of the molt, by controlling the JH titer. Third, this study demonstrates that EPPAT at proper doses is a viable probe for studying enzyme and hormone action in vivo without pharmacological artifacts.

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Résumé Cette étude indique d'abord que, lorsque la dégradation de l'hormone juvénile (JH) par la JHE est bloquée par un inhibiteur (EPPAT, O-éthyl-S-phényl-phosphoramidothiolate) les effets prothoracicotropiques—libération d'ecdysone—sont retardés chez Trichoplusia ni Hübner. Ainsi, la JHE est un élément important de la dégradation de JH, impliquant que sans une dégradation régulière, la teneur en JH deviendra anormalement élevée. Cette accumulation d'hormone juvénile endogène chez les larves traitées à l'EPPAT provoque de plus une mue larvaire supplémentaire. Par conséquent, JHE est importante aussi dans le contrôle de la nature de la mue, en déterminant la teneur en JH. Enfin, cette étude a montré que l'EPPAT à des doses appropriées est un moyen efficace pur étudier l'action hormonale in vivo sans artéfacts pharmaceutiques.

     

Degradation of a radiolabeled juvenile hormone analog using two insect species

A synthetic insect juvenile hormone analog (a juvenoid), ethylN-[2-[4-[[2,2-(ethylenedioxy)cyclohexyl]methyl]phenox]ethyl]carbamate, which has displayed high biological activity against different insect species and high stability under field conditions, was selected as a biologically active model compound for a study of a juvenile hormone analog degradation. The biologically active compound itself and its three diversely radiolabeled derivatives were applied to the flesh fly (Sarcophaga bullata) or the tsetse fly (Glossina palpalis), respectively. Monitoring of a fate of the applied juvenile hormone analog was carried out using a detection method of the radioactivity microdistribution within the whole insect body in combination with a radio high performance liquid chromatography (radio-HPLC), both of whole-body extracts made in different, but in advance scheduled, time intervals, and of extracts of insect excreta accumulated over an eight-day experiment.     

Somatic embryogenesis in polyembryonic Secale cereale L.

Summary The progeny of polyembryonic Secale cereale L., was used to study the in vitro response of the immature embryos. The formation of embryogenic calli was very high, and this response and its distribution was statistically different to that shown by the normal regenerated plants and the original population. This behaviour seems to be related to a genetic condition which favours the presence of supernumerary embryos, in vivo as well as in vitro.     

Juvenile hormone effects on polymorphism in the pea aphid, Acyrthosiphon pisum

When reared in short days (LD 12:12) at 15°C, apterous Acyrthosiphon pisum gave birth to sexual females (oviparae) exclusively for the first eight days of larviposition. After this time they switched to the production of parthenogenetic females (viviparae). Topical application of juvenile hormones I, II and III to fourth instar or adult ovipara-producers induced the precocious appearance of parthenogenetic females in the progeny sequence. Various forms intermediate between oviparae and viviparae were also produced and repetitive JH-I treatments resulted in a few alatiform progeny. However, many of the JH induced apterous, parthenogenetic females appeared to be normal viviparae and were capble of reproduction. Thus, prenatal treatment of oviparous embryos with JH diverts development towards the viviparous form. JH-I treatment of long-day reared A. pisum had no effect on the type of progeny produced.

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Effets de l'hormone juvénile sur le polymorphisme d'Acythosiphon pisum

Résumé Quand il est élevé sous jours courts (LN 12/12) à 15°C, le type anglais vert d'Acyrthosiphon pisum ne donne naissance qu'à des femelles sexuées (ovipares) pendant la première partie de sa période de reproduction. Ensuite quelques types intermédiaires ovipares/vivipares peuvent apparaitre avant que les pucerons ne bifurquent spontanément vers la production de femelles parthénogénétiques (vivipares). L'application cutanée d'hormones juvéniles (JH I, II, et III) aux larves de quatrième stade ou à des adultes producteurs d'ovipares provoque l'apparition prématurée d'intermédiaires et de vivipares dans la descendance. Les différentes formes intermédiaires produites par des applications répétées de J.H. comprenaient des types ailés ou partiellement ailés. Cependant, les vivipares aptères induits par J.H. étaient morphologiquement normaux et beaucoup étaient capables de se reproduire. Des traitements semblables aux J.H. de vivipares élevées en jours longs (LN 16/8) n'ont pas eu d'effets sur le type de la descendance.On ne sait pas si l'action de JH exogène sur l'induction des vivipares est direct ou indirect. La reprogrammation des embryons, autrement destinés à se développer comme ovipares, est examinée en relation avec notre connaissance du contrôle endocrine du polymorphisme des pucerons.

     

In vitro induced pinocytotic activity by a juvenile hormone analogue in oocytes of Drosophila melanogaster

Summary Pinocytotic activity has been analyzed in Drosophila oocytes following either in vivo or in vitro exposure to horseradish peroxidase. The enzyme tracer gains access to the yolk spheres only when supplied to the oocyte in vivo. In oocytes cultured in vitro, peroxidase remains restricted to the residual coated vesicles and to the tubular profiles formed in excess in the cortical ooplasm.In an attempt to induce peroxidase uptake by oocytes cultured in vitro, various incubations were tested. Among these, hemolymph from both sexes is capable of promoting peroxidase uptake up to a level comparable to that detectable in vivo. On the other hand, fat body extracts fail to promote such cellular activity. Finally, the juvenile hormone analogue ZR-515 is shown to be the only factor required to promote pinocytotic activity under the experimental conditions tested. The observations are interpreted to indicate that vitellogenin has no inductive role on pinocytosis but simply acts by adhering to the forming coated vesicles which in turn are produced by the oolemma in response to the action of juvenile hormone.     

Juvadecene: Discovery of a juvenile hormone mimic in the plant,Macropiper excelsum

A compound with significant insect juvenile hormone activity was isolated from the plant, Macropiper excelsum. The chemical structure was determined by spectral methods to be 1-(3,4-methylenedioxyphenyl)-trans-3-decene(l), and confirmed by synthesis. The hormonally active substance applied topically to last (fifth)-instar nymphs of the milkweed bug (Oncopeltus fasciatus) induced a supernumerary metamorphosis at 30 μg. Higher doses were toxic.     

Effects of methoprene and juvenile hormone on larval ecdysis,emergence, and metamorphosis of the endoparasitic wasp, Apanteles congregatus

Topical application of juvenile hormone I and III or the hormone analogue methoprene to parasitized Manduca sexta larvae inhibited subsequent emergence of the endoparasitic wasp Apanteles congregatus. Methoprene treatment inhibited wasp emergence in a dose-dependent manner, causing either a delay or total inhibition of emergence. These results were interpreted as reflecting inhibitory effects of juvenile hormone on the second-larval ecdysis of the parasitoid that normally occurs during emergence from the host larva. Parasitoid ecdysis was disrupted even when methoprene was applied to host larvae a few hours prior to the normal expected time of emergence. A correlation between the number of emerging parasitoids and the timing of emergence was seen in methoprene-treated hosts, and few parasitoids emerged after day 9 of the host's fifth-instar. Our findings suggest that the suppression of emergence by juvenile hormone analogues noted in previous studies may be due to a similar inhibitory effect on parasitoid ecdysis. We also observed that parasitoids emerging from hosts treated with a low dose of methoprene (1 μg) later pupated normally but then formed nonviable pupal-adult intermediates. Thus use of this insect growth regulator must be undertaken carefully to prevent possible adverse effects on natural parasitoid populations.     

Physiological effects of a juvenile hormone analog on larvae of Spodoptera littoralis

Topical application of JHA to Spodoptera littoralis larvae extends the duration and increases the final weight of the last larval instar. No supernumary moults occur. Respiration decreases to very low levels and glycogen and lipid stores increase. Respiration remains lower in JHA-treated pharate pupae and more polyunsaturated C₁₈ fatty acids accumulate than in normal pharate pupae. These physiological parameters are similar to those occurring during diapause but are not definitive. Diapause has not been reported in field populations of S. littoralis, nor it's artificial induction by photoperiodic manipulation.

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Résumé Le dernier stade des chenilles de Spodoptera littoralis, traitées avec JHA, est nettement prolongé par rapport aux témoins. Bien que le poids final des chenilles traitées soit plus élevé que celui des témoins, il n'y a pas de mue surnuméraire. Ces chenilles géantes s'alimentent rarement et leur respiration est sérieusement réduite. Les réserves de glycogène s'accumulent chez les chenilles traitées à la même date chronologique que chez les prénymphes non traitées. Bien que les prénymphes traitées à la JHA conservent leur profil biochimique juvénile — tout au moins dans la mesure où on considère la teneur en glycogène et en lipides totales — leur composition en acides gras saturés et non saturés diffère de celle normalement rencontrée à la fin du développement larvaire, et beaucoup plus d'acide oléique monosaturé s'accumule finalement chez les insectes traités par JHA. Ces résultats suggèrent que la JHA fournie peut avoir provoqué un état de développement de diapause naissante semblable par quelques traits fondamentaux à celle induite par la JH endogène chez les insectes enclins à la diapause. Il est cependant prématuré de définir cet état physiologique comme indiquant une vraie diapause chez Spodoptera littoralis.

     

The effect of farnesyl methyl ether, a mimic of insect juvenile hormone, on Hymenolepis Diminuta in vitro

The effect of farnesyl methyl ether, a mimic of insect juvenile hormone, on Hymenolepis diminuta in vitro. International journal for Parasitology4: 211–218. The inhibition of weight gain of Hymenolepis diminuta in vitro by farnesyl methyl ether (FME) was not dose-dependent, so that there were no differences produced by FME in concentrations from 10^?5 to 10^?13m. At all concentrations of FME used the weight gain over a 6-day culture period was half of that of controls. The FME-treated and control worms were identical to each other in proglottid number and the degree of maturity. Neither the dry to wet weight ratios of worms nor the carbohydrate, protein or lipid ratios were altered by FME. The release of neurosecretory material from the neurosecretory cells in the rostellum (as measured by the degree of their fuchsinophilia) was more advanced in time in the FME-treated worms than in the controls, and it is suggested that this premature release of neurosecretion, triggered either directly or indirectly by traces of FME in the medium, upset a control mechanism in the germinative tissue of the neck region, which determines the mass, but not the number, of the proglottids in the strobila.     

Presence of soldier larvae determines the outcome of competition in a polyembryonic wasp

Soldier‐producing polyembryonic waSPS are the only social animals that develop as parasites inside the bodies of other insects. Characterizing the kin composition of broods is central to understanding the evolution of the soldier caste in these unique social insects. Here we studied the role of soldiers in mediating the outcome of competition among clones of the polyembryonic wasp Copidosoma floridanum. Soldier‐producing female clones usually monopolized host resources, whereas soldierless male clones usually coexisted in hosts. Behavioural experiments further indicated that early‐emerging soldiers are specialized to combat intraspecific competitors and later‐emerging soldiers are specialized for defence against interspecific competitors. Taken together, our results point to intraspecific competition as a major selective force in the evolution of the soldier caste. Our data also present an evolutionary conundrum: given the benefit of soldiers, why are male clones functionally soldierless?     

Metabolism of juvenile hormone in cultures of ovaries and fat body in the cockroachperiplaneta Americana

Summary The time course of juvenile hormone (JH) metabolism is examined in cultures ofPeriplaneta americana fat body and ovaries in medium containingManduca sexta carrier protein or cockroach hemolymph. In the absence ofM. sexta carrier protein or cockroach hemolymph, both tissues extensively catabolize exogenous [³H]JH in the medium. Addition of the carrier protein or hemolymph to the culture system prevents the hydrolysis of the hormone in the medium. Within the tissues JH is degraded whether or not carrier protein or hemolymph is present which suggests that the protective role of these molecules is exclusively extracellular. Incubation of [³H]JH with medium preconditioned with tissue results in destruction of the hormone. This suggests that the fat body secretes esterases into the medium. In contrast, the ovarioles hydrolyze the hormone by means of cell-associated enzyme. The relationship of these phenomena to insect development is discussed. This work supported by NSF Grant PCM 76-02229 and University of Kansas Biomedical Sciences Grant RR-07037.     

Effect of Bt-cotton expressing Cry1A(c) on the survival and fecundity of two hymenopteran parasitoids (Braconidae, Encyrtidae) in the laboratory

We examined the effect of Bt-cotton (Event 531) plants expressing the Bacillus thuringiensis δ-endotoxin CryIA(c) on two hymenopteran endoparasitoids, Cotesia marginiventris and Copidosoma floridanum. In the laboratory, parasitized and unparasitized Pseudoplusia includens larvae were reared on foliage from a conventional soybean cultivar (Pioneer 97B61), a conventional cotton cultivar (DPL 5415), or a Bt-cotton cultivar (NuCotn 33B). C. marginiventris developed significantly faster within P. includens larvae feeding on Pioneer 97B61 and DPL 5415 compared to those feeding on NuCotn 33B. C. marginiventris that developed inside P. includens larvae feeding on NuCotn 33B suffered reduced longevity, and females had fewer ova. NuCotn 33B also affected the growth and development of P. includens parasitized with C. floridanum and life history parameters of adult C. floridanum. Parasitized and unparasitized P. includens developed more slowly when they were fed NuCotn 33B and the prepupae weighed less. Survival of parasitized and unparasitized P. includens was lower when larvae were fed NuCotn 33B and some evidence points to higher susceptibility of parasitized caterpillars to intoxication by NuCotn 33B. Fewer C. floridanum adults emerged from hosts fed NuCotn 33B, but pupal weight and adult longevity were unaffected. Analysis comparing the two experiments conducted with C. floridanum suggests that older NuCotn 33B plants (90–120 days after planting) may affect parasitoid development and adult survival less than younger NuCotn 33B plants (60–90 days after planting). Feeding on NuCotn 33B by P. includens affected the survival and development of the two hymenopteran endoparasitoids studied here, and the degree of the effect was similar to that observed with natural resistance found in soybean plants. It remains to be determined if the effects demonstrated here are less than, equal to, or greater than the impact of conventional insecticide applications used in conventional, non-transgenic, cotton.     

Effects of the ecdysteroid agonists RH 5849 an RH 5992, alone and in combination with a juvenile hormone analogue, pyriproxyfen, on larvae ofSpodoptera exigua

Biological activity assays with RH 5849 and RH 5992 indicated that both compounds affected growth and development of last-instar larvae ofSpodoptera exigua (Hübner) (Lepidoptera: Noctuidae) in a dose-dependent manner. Within the first 24 h after treatment by continuously offering leaves dipped in a water solution of ≥50 mg/l RH 5849 and ≥0.5 mg/l RH 5992, symptoms of a prematurely induced larval moult and head capsule apolysis were visible. Intoxicated larvae died shortly afterwards, showing signs of unsuccessful ecdysis. LC₅₀-values of RH 5849 and RH 5992 for fifth-instarS. exigua larvae were 110 and 2.5 mg/l, respectively. Pyriproxyfen alone affected the larval stage and disturbed normal metamorphosis. One supernumerary larval instar occurred occasionally. LC₅₀-value for pyriproxyfen was 1.7 mg/l. Larvae simultaneously treated with RH 5849 or RH 5992 and pyriproxyfen, continued to grow until they attained a size and weight about 2–3 times that of the controls. This growth was accompanied by at least one and sometimes two supernumerary moults. Concerning thein vivo imaginal wing disc growth and development, only in larvae treated with 10 and 50 mg/l RH 5849 or 0.5 mg/l RH 5992, tracheole migration was observed earlier than in the controls. When applying 300 mg/l RH 5849 or 3–7 mg/l RH 5992, the discs remained small and no signs of tracheole migration were observed. In larvae simultaneously treated with RH 5849 or RH 5992 and pyriproxyfen, tracheole migration was not prematurely induced and a pupal cuticle was produced in the discs of larvae, undergoing a supernumerary moult. No clear signs of evagination were observed.