Temporal change in the location of egg-laying by a bud-galling sawfly,Euura mucronata,on growing shoots of Salix cinerea

Summary Temporal change in the egg-laying behaviour of Euura mucronata was studied on growing shoots of Salix cinerea. Many new shoots arise from each previous year's shoot of S. cinerea; they grow very rapidly at the beginning of the oviposition period of E. mucronata, but their growth pattern changes dramatically during the 3 weeks of oviposition. When oviposition commences, all new shoots are equally long, and E. mucronata females lay eggs on them randomly; but toward the end of oviposition, eggs are laid on the longest of the new shoots, which originate on the distal part of the previous year's shoot. By the end of oviposition, the mean proportion of ovipositor borings that had resulted in egg-laying was 65.8% on distal shoots and declined linearly to 25% on shoots arising from the base of the previous year's shoot. The proportion of attacked buds from which larvae emerged declined non-linearly from 13.8% on the distal shoot to 3.5% on the basal shoots. The position of the shoot accounted for 67% and 73% of the variance in success of egg-laying and larval development, respectively.     

Incidence of apple fruit and leaf surface metabolites on Cydia pomonella oviposition

Water soluble metabolites identified from surfaces of apple tree fruit and leaf stimulate oviposition in Cydia pomonella L. The effects of two artificial blends of primary metabolites representing fruit and leaf surfaces, respectively, and of components within the blends were examined on egg-laying after two time periods: 3 min and 25 min of darkness. An artificial mixture of six metabolites, viz., three sugar alcohols (sorbitol, quebrachitol, and myo-inositol) and three sugars (glucose, fructose, and sucrose) did stimulate egg-laying. Fructose, sorbitol, and myo-inositol are important components of the stimulatory blend. Contact durations may induce variations in egg-laying responses. After 3 min of darkness, there were no differences in numbers of females laying eggs nor in the numbers of eggs laid on cloths treated with the complete blends and the controls. There were, however, clear effects of groups of compounds and of individual compounds. Reduced blends without sugars and sugar alcohols were in many cases significantly less stimulatory than the complete blends and the controls. After 25 min of darkness, the proportions of females laying eggs as well as the numbers of eggs were higher after treatment with the complete blends than on the controls. The proportions of females laying eggs on cloths treated with the reduced blends were rather similar to the controls, whereas there were still significant effects on the numbers of eggs laid after treatment with reduced blends derived from fruits but not from leaves.     

Effects of time-dependent intraspecific competition on offspring survival in the butterfly, Anthocharis scolymus (L.) (Lepidoptera: Pieridae)

Effects of mothers' eclosion and oviposition timing on the survival of their offspring in the pierid butterfly Anthocharis scolymus (L.) was examined. I recorded the performance of individual eggs and larvae that differed in their mother's eclosion and oviposition timing in a natural population, where A. scolymus feeds on Turritis glabra (L.) Bernh. Eggs laid early in the season, and larvae emerging from these eggs, had higher survival than eggs laid later, and larvae emerging from eggs that were laid later in the season, the causal factor being egg cannibalism by larvae on the same host plant. Logistic regression showed that females eclosing early in the season had higher offspring survival than females eclosing later. I conclude that optimal timing of adult eclosion in A. scolymus is a trade-off between eclosing early with associated higher offspring survival but lower egg-laying rate, and eclosing later with associated lower offspring survival but higher egg-laying rate. Received: 4 August 1997 / Accepted: 19 October 1997     

A circuit model of the temporal pattern generator of <Emphasis Type="Italic">Caenorhabditis</Emphasis> egg-laying behavior

Background  

Egg-laying behavior in the nematode C. elegans displays a distinct clustered temporal pattern: egg-laying events occur primarily in bursts or active phases, separated by inactive phases during which eggs are retained. The onset of the active phase can be modeled as a Poisson process with a time constant of approximately 20 minutes, while egg-laying events within an active phase occur with a faster time constant of approximately 20 seconds. Here we propose a cellular model for how the temporal pattern of egg-laying might be generated, based on genetic and cell-biological experiments and statistical analyses.     

Factors affecting egg predation in Black-tailed Gulls

In colonial seabirds, nesting density, egg-laying date and nest microhabitat affect the probability of eggs being taken by avian predators. Jungle Crows (Corvus macrorhynchos) are dominant predators of eggs of Black-tailed Gulls (Larus crassirostris). Factors affecting the probability of gulls allowing the crows to attack their nests or depredate their eggs and the probability of eggs being taken were studied by direct observation and egg census, respectively. The effect of vegetation heights, position in the colony, egg-laying date and neighbour nests on the probability of eggs being taken were examined at multiple spatial scales. Gull nests were depredated more easily by larger groups of crows. Nests in peripheral areas (<4 m from the edge of the colony) were also depredated more easily by the crows walking on the ground. Although the nests where eggs were laid early in the season were depredated more frequently, such nests highly synchronised in egg laying within a <2-m radius were less likely to be depredated than less-synchronised nests. The nests in tall vegetation were less likely to be depredated though those having neighbour nests in tall vegetation were not. The number of neighbour nests did not affect the probability of eggs being taken. Antipredation effects of nesting microhabitats vary with spatial scales at which the crows search and attack the nests of gulls.     

Egg-laying experience and acceptance of parasitized hosts by the parasitoid,Leptopilina heterotoma (Hymenoptera: Eucoilidae)

The influence of egg-laying experience on the response of females of the eucoilid parasitoid,Leptopilina heterotoma, to parasitized and unparasitizedDrosophila melanogaster host larvae was examined under more controlled conditions than those used in past studies. In laboratory assays, we precisely manipulated both the number of eggs laid by females and the kind of larvae (parasitized versus unparasitized) in which the eggs were laid. We found that the tendency to avoid laying eggs in parasitized hosts depended markedly on whether or not eggs had been laid previously, but depended little on whether those eggs had been laid in parasitized or unparasitized hosts. The observed effect of general egg-laying experience on avoidance of parasitized hosts may reflect responses to either changes in the wasp's internal state (perhaps, changes in egg load) or changes in the wasp's neural representation of the external environment (such as those presumed to occur during learning). In light of these results, we offer a tentative reinterpretation of several earlier studies.     

Why walnut flies superparasitize: time savings as a possible explanation

 This study evaluated a possible fitness advantage, specifically time savings, that might account for an unusual propensity in walnut flies (Rhagoletis spp.) to superparasitize their walnut hosts and to place eggs into existing egg-laying cavities. The first part of this study demonstrated that, in laboratory assays, females of two walnut fly species, R. boycei and R. juglandis, save time when cavities are reused and that in R. juglandis, where it was examined in detail with in vivo staining of eggs, time saving was not an artifact of differences in the size of clutches deposited at new versus existing sites. We further demonstrated that time savings reflected a reduction in the time required to generate the cavity itself. In the second part of the study, we evaluated the possibility that, in the field, time saved by reusing existing cavities is nullified by extra time spent mating associated with a previously described tendency for males to guard these cavities. Field observations of R. juglandis indicated that use of existing sites was, as expected, associated with increased mating. Yet, despite the added time spent mating, in observations of similar length females attempting to lay eggs at existing sites deposited clutches more often than females attempting to lay eggs at new sites. We discuss these results in the context of the more common pattern of superparasitism avoidance observed in host-specific insects. Received: 25 January 1996 / Accepted: 25 June 1996     

Life-history variation in selfing multilocus genotypes of the land slug Deroceras laeve (Pulmonata: Agriolimacidae)

The land slug Deroceras laeve shows an enormous variation inlife-history traits. Here, we compared several life-historycharacters (i.e. length of the prereproductive period, growthduring the egg-laying period, number of eggs, hatching success)within and among five selfing homozygous multilocus genotypes(MLG) and investigated a possible trade-off between the numberof eggs produced during the egg-laying period and growth duringthe egg-laying period. Life-history traits were highly variable.There was no significant correlation between slug size and fecundity.Slugs that matured early produced more eggs than slugs thatmatured later. Life-history traits were significantly differentamong MLGs, suggesting that genetic factors may affect and explainat least part of the variation in life-history traits in D.laeve. Finally, we found no evidence of a trade-off betweenthe number of eggs produced during the egg-laying period andgrowth during the egg-laying period. (Received 1 June 2005; accepted 28 October 2005)     

Laboratory rearing of the squirrel fleaCeratophyllus sciurorum sciurorum with notes on its biology

A rearing method for the fleaCeratophyllus (Monopsyllus) sciurorum sciurorum, a pest on farmed mink in Europe, has been developed. The behaviour of egg-laying and the development period are described at 23 °C and 80% r.h. The eggs ofC. sciurorum sciurorum hatched after 4–7 days and cocoons were formed 10–12 days later. The development period from egg to newly emerged adult was 23–32 days. Females emerged, in general, earlier than males.     

Pear rust mite,Epitrimerus pyri (Acari: Eriophyidae) oviposition and nymphal development on Pyrus and non-Pyrus hosts

The ovipositional response of deutogynes of the pear rust mite, Epitrimerus pyri Nalepa, and the rate of development of E. pyri nymphs differed according to the host plant to which mites were exposed. Among the fourteen Pyrus hosts examined, leaves from the Clapp's Favorite cultivar of P. communis elicited the strongest egg-laying response and the fastest rate of nymphal development, whereas egg-laying was least and nymphal development slowest on leaves from P. calleryana. Among the three non-Pyrus hosts, E. pyri oviposited and completed nymphal development on apple and quince leaves, but would not deposit eggs on apricot. Antixenosis appears to be the primary mechanism of resistance, although antibiosis may be operating to a lesser extent. These results are discussed in relation to the resistance of Pyrus to other arthropod pests.     

Rhythmic egg-laying behaviour in virgin females of fruit flies Drosophila melanogaster

Fruit fly Drosophila melanogaster females display rhythmic egg-laying under 12:12?h light/dark (LD) cycles which persists with near 24?h periodicity under constant darkness (DD). We have shown previously that persistence of this rhythm does not require the neurons expressing pigment dispersing factor (PDF), thought to be the canonical circadian pacemakers, and proposed that it could be controlled by peripheral clocks or regulated/triggered by the act of mating. We assayed egg-laying behaviour of wild-type Canton S (CS) females under LD, DD and constant light (LL) conditions in three different physiological states; as virgins, as females allowed to mate with males for 1?day and as females allowed to mate for the entire duration of the assay. Here, we report the presence of a circadian rhythm in egg-laying in virgin D. melanogaster females. We also found that egg-laying behaviour of 70 and 90% females from all the three male presence/absence protocols follows circadian rhythmicity under DD and LL, with periods ranging between 18 and 30?h. The egg-laying rhythm of all virgin females synchronized to LD cycles with a peak occurring soon after lights-off. The rhythm in virgins was remarkably robust with maximum number of eggs deposited immediately after lights-off in contrast to mated females which show higher egg-laying during the day. These results suggest that the egg-laying rhythm of D. melanogaster is endogenously driven and is neither regulated nor triggered by the act of mating; instead, the presence of males results in reduction in entrainment to LD cycles.     

Host-plant susceptibility to the carrot fly, Psila rosae. 1. Acceptability of various host species to ovipositing females

A preference hierarchy of the oligophagous carrot fly for 30 umbelliferous host-plant species or varieties and six non-hosts was assessed. Foliage of the test plants was presented to a laboratory population of the fly in oviposition choice experiments together with leaves of a standard plant, the susceptible carrot cultivar “Danvers”. Only two species (Anthriscus cerefolium, Carum carvi) were more acceptable than the standard plant, while about half the species received significantly fewer eggs. Some umbellifers (Anethum graveolens, Pimpinella major) had a low acceptability similar to that of non-hosts (non-umbelliferous plants). The results obtained with dual and multiple choice assays were in agreement. A multiple choice assay with potted plants yielded a similar ranking of the species as assays using detached leaves, indicating that foliage is representative for whole plants. Variability in the exploratory runs performed by the females on the leaves prior to egg-laying is described and quantified for hosts and non-hosts. Differences in post-alighting pre-ovipositional behaviour of the female flies on the leaves accounted for a major part of the variation recorded in egg-laying.     

L'élevage massif artificiel deCeratitis capitata Wied.

Summary A method of artificial mass-breeding ofCeratitis capitata has been developped after the results obtained byFinney andChristenson. The breeding of the larvae is obtained on a medium made of powdered atomised carrot and brewer's yeast, the development of molds and bacteria being inhibited by benzo?c and hydrochloric acids. Nearly 80 per cent of the eggs give adult flies. An original method for obtaining the eggs in an artificial egg-laying apparatus is described. The development of the insect is completed in 21 days at the temperature of 26°C. A daily production of 20 000 flies or more is easily obtained, at very low cost, by the work of a single person.

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Travail réalisé avec la collaboration technique de MlleM. Barthes.

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Station de Zoologie agricole du Centre de recherches agronomiques du Sud-Est.     

Abnormal egg-laying behaviour of female carrot flies (Psila rosae) induced by the fungus Entomophthora muscae

Female carrot flies (Psila rosae F.) infected with Entomophthora muscae (C.) Fres. on days 1 to 4 of adult life were unable to lay their eggs near the food plant. Flies infected on day 5 of adult life showed an egg-laying behaviour similar to that of uninfected flies. E. muscae-infected female carrot flies caught in the field showed an egg-laying behaviour comparable to that of the flies infected in the laboratory during the first day of their live.Eggs laid by infected females were equally as fertile as eggs laid by uninfected females, and developed into normal flies.It is concluded that female carrot flies infected with E. muscae in the field do not in general contribute to the development of the carrot fly population, due to their abnormal egg-laying behaviour.

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Zusammenfassung Wenn Weibchen der echten Möhrenfliegen (Psila rosae) zwischen den ersten und der vierten Tag in dem Laboratorium mit E. muscae infiziert worden waren, könnten sie ihre Eier an den Futterpflanze nicht ablegen. E. muscae-infizierte Weibchen, die ins Feld gefangen worden waren, zeigten auch ein abweichendes Eiablageverhalten. Ihr Behalten war mit dem Verhalten von Fliegen vergleichbar, die an ihrem ersten Lebenstag im Laboratorium infiziert worden waren.Von infizierten Weibchen gelegte Eier waren ebenso fruchtbar wie die von nicht infizierten Weibchen gelegten Eiern. Es wird konkludiert, dass E. muscae-infizierten Weibchen der echten Möhrenfliegen der Entwicklung der Population keinen Beitrag leisten.

     

Factors promoting maternal trophic egg provisioning in non-eusocial animals

The adaptive function of trophic egg-laying is generally regarded as extended parental investment to the offspring. However, the evolutionary factors promoting trophic egg-laying are still unclear, because the amount of maternal investment per offspring should be ideally equal between smaller offspring with trophic eggs and larger offspring without any additional investment. Several authors have suggested that trophic egg-laying should evolve only when egg size is constrained, but this hypothesis has not been evaluated. We investigated the evolutionary mechanisms of trophic egg-laying by two different approaches. First, we evaluated morphological constraints on egg size in two sibling ladybird species, Harmonia axyridis, which is known to produce trophic eggs, and H. yedoensis. Second, we theoretically predicted the optimal proportion of trophic eggs to total eggs and egg size in relation to environmental heterogeneity, predictability of environmental quality, and investment efficiency of trophic egg consumption. The intra- and interspecific morphological comparisons suggest that morphological constraints on the evolutionary determination of egg size are weak at best in the two ladybird species. Moreover, we theoretically showed that small egg size and trophic egg-laying are favoured in heterogeneous environments when mothers cannot adjust egg size plastically. We also showed that even a small reduction in investment efficiency makes a trophic egg strategy unlikely, despite relatively high environmental predictability. We conclude that trophic egg provisioning may be a flexible maternal adaptation to a highly heterogeneous environment rather than a response to a morphological constraint on egg size.     

Copulation, egg production and sex-ratio in Phytoseiulus persimilis and Amblyseius bibens (Acari: Phytoseiidae)

In Phytoseiulus persimilis Athias-Henriot and Amblyseius bibens Blommers a relation was found between the duration of copulation and the resultant egg production. After 15 min of copulation no eggs were produced but after respectively 30, 60 and 90 min an increasing number of eggs were deposited. In P. persimilis the total number of eggs can be produced after one copulation of long duration but in A. bibens several copulations are necessary for total egg production. Evidence was found that egg deposition depends on the quantity of sperm transferred during insemination. Relatively more males developed from eggs deposited during the first day of egg deposition but males are produced throughout the whole egg-laying period. Females of P. persimilis remain fertile throughout their life until they have deposited their total possible number of eggs. Three to 4 egg-laying periods could be induced by copulations of 30 min duration. At each period relatively more males developed from eggs laid during the first days. Males of P. persimilis were found to be very fertile. The results obtained are discussed in relation to the reproductive strategy of both species.

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Zusammenfassung Bei Phytoseiulus persimilis Athias-Henriot und Amblyseius bibens Blommers wurde ein Zusammenhang gefunden zwischen Kopulationsdauer und der resultierenden Eiproduktion. Nach 15 Minuten Kopulation wurden keine Eier produziert; nach 30, 60 und 90 Minute Kopulation hingegen wurde eine steigende Zahl Eier abgelegt. Bei P. persimilis kann die totale Eizahl nach einer einzigen Kopulation von langer Dauer produziert werden, während bei A. bibens mehrere Kopulationen notwendig sind. Es wurden Anhaltspunkte gefunden, dass die Eizahl von der während der Besamung übertragenen Spermienmenge abhängt. Aus während der ersten Tage der Eiproduktion abgelegten Eiern entwickelten sich relativ mehr Männchen, doch wurden Männchen während der ganzen Eiablageperiode produziert. Weibchen von P. persimilis bleiben während ihres ganzen Lebens, bis sie alle Eier abgelegt haben, fertil. Drei bis vier Eiablageperioden konnten durch Kopulationen von 30 Minuten Dauer induziert werden. Bei jeder Periode entwickelten sich aus den in den ersten Tagen abgelegten Eiern relativ mehr Männchen. Männchen von P. persimilis erwiesen sich als sehr fertil. Die Resultate werden im Zusammenhang mit der Vermehrungsstrategie beider Arten diskutiert.

     

A novel procedure for obtaining denuded sea urchin eggs and observations on the role of the vitelline layer in sperm reception and egg activation

A procedure is described for the complete removal of the vitelline layer of the eggs of the sea urchin, Strongylocentrotus purpuratus. The method involves treatment of unfertilized eggs with an S. purpuratus cortical granule protease preparation followed by incubation in an alkaline dithiothreitol seawater solution. Eggs denuded of their vitelline layers react metabolically to parthenogenetic agents and sperm like unfertilized eggs, whereas the fertilizability of denuded eggs and receptivity to sperm is much less than controls. The present method is superior to previous methods using mercaptans in that all of the vitelline layer is removed and to procedures using other proteolytic enzymes in that no ¹²⁵I-labelled plasma membrane proteins are extensively modified. Thus the cortical granule protease dithiothreitol procedure is ideal for studies of the plasma membrane of the unfertilized egg and for studies on the role of the vitelline layer in normal fertilization and development.     

Measuring the Effects of Bacteria on C. Elegans Behavior Using an Egg Retention Assay

C. elegans egg-laying behavior is affected by environmental cues such as osmolarity¹ and vibration². In the total absence of food C. elegans also cease egg-laying and retain fertilized eggs in their uterus³. However, the effect of different sources of food, especially pathogenic bacteria and particularly Enterococcus faecalis, on egg-laying behavior is not well characterized. The egg-in-worm (EIW) assay is a useful tool to quantify the effects of different types of bacteria, in this case E. faecalis, on egg- laying behavior.EIW assays involve counting the number of eggs retained in the uterus of C. elegans⁴. The EIW assay involves bleaching staged, gravid adult C. elegans to remove the cuticle and separate the retained eggs from the animal. Prior to bleaching, worms are exposed to bacteria (or any type of environmental cue) for a fixed period of time. After bleaching, one is very easily able to count the number of eggs retained inside the uterus of the worms. In this assay, a quantifiable increase in egg retention after E. faecalis exposure can be easily measured. The EIW assay is a behavioral assay that may be used to screen for potentially pathogenic bacteria or the presence of environmental toxins. In addition, the EIW assay may be a tool to screen for drugs that affect neurotransmitter signaling since egg-laying behavior is modulated by neurotransmitters such as serotonin and acetylcholine^5-9.     

Self-restraint and sterility in workers of Acromyrmex and Atta leafcutter ants

Summary. Queens of leafcutter ants (Acromyrmex and Atta) are highly multiply mated, resulting in a potential queenworker and worker-worker conflict over who should produce the males in the colony. We studied whether this conflict is expressed, by determining the amount of reproductive egg-laying by workers in queenright colonies of Acromyrmex echinatior, Acromyrmex octospinosus, Atta cephalotes, and Atta sexdens through ovary dissections. Worker sons are absent or rare in queenright Acromyrmex colonies, but can be produced in orphaned colonies. In Atta, most workers have rudimentary ovaries that never produce eggs, but a few (mostly small and medium workers that form a retinue around the queen) lay many trophic eggs for consumption by the queen. These eggs are large, flaccid, and lacking in yolk compared to queen-laid eggs, and appear to be always inviable. In Acromyrmex, many workers (especially young large workers) lay eggs that are similar in size to queen-laid eggs, but mostly with a reduced amount of yolk. Trophic eggs appear to be an important source of food for larvae in Acromyrmex (especially in Ac. echinatior), but not in Atta. Five (0.8) of 616 dissected Ac. echinatior workers but no Ac. octospinosus workers (n = 552), had ready-to-lay reproductive eggs. Old workers in all four species are incapable of laying eggs due to ovary resorption. We conclude that Atta workers are sterile, while Acromyrmex workers display reproductive self-restraint, possibly representing an earlier stage in the evolution towards worker sterility. Worker selfrestraint in Acromyrmex may be maintained by a queen or worker policing mechanism, but individual cost-benefit explanations may also apply.Received 1 March 2004; revised 28 June 2004; accepted 1 July 2004.     

BREEDING BIOLOGY AND BEHAVIOUR OF THE QUAIL FINCH ORTYGOSPIZA ATRICOLUS

Summary

Nuttall, R.J. 1992. Breeding biology and behaviour of the Quail Finch Ortygospiza atricollis. Ostrich 63:110-117.

During a study of the breeding biology of the Quail Finch Ortygospiza atricollis, observations of nest-building, egg-laying, incubation and nestling periods, and nestling development in a grassland near Pietermaritzburg, South Africa were supplemented with observations of breeding behaviour in captivity. Mean clutch size was 4,5 and eggs were laid at intervals of approximately one day. Incubation began after the third or fourth egg was laid. An incubation period of 15–16 days and an estimated nestling period of 18–19 days was recorded. Incubation and brooding are shared by both sexes. Breeding success was low (26,7% ?28,6%), with most losses resulting from predation during either the egg-laying or incubation stages.