The maturation divisions of the parthenogenetic stick insect Carausius morosus Br. (Orthoptera,Phasmidae)

The meiotic stages in the maturation of the egg of the parthenogenetic stick insect Carausius morosus Br. is described. The meiosis consists of two equational divisions and parthenogenesis is thus apomictic. The first prometaphase is formed between 5.8–5.5 days before oviposition; the first metaphase lasts until oviposition; the meiosis ends between 14 and 24 hours after oviposition. An extra chromosome doubling occurs before first anaphase, causing the first metaphase chromosomes to resemble bivalents and requiring that meiosis consists of two divisions instead of one. A terminal affinity between the daughter chromosomes influences chromosome movement during first and second metaphase and anaphase. The first and second polar bodies degenerate. The first polar body divides amitotically during pycnosis. Meiosis takes place ventrally in the egg, the germ anlage development dorsally. The pronucleus divides mitotically in the ventral part of the egg during its migration from ventral to dorsal, enabling blastoderm development to take place both ventrally and dorsally.     

Effects of 2,4-D and maleic hydrazide on the glycogen level in the embryonic development of Pimpla turionellae (L.) (Hym., Ichneumonidae)

Abstract: The effects of the herbicides 2,4-D and maleic hydrazide, which are used against wild plants in farming, on the glycogen level in the embryological development of Pimpla turionellae, an important insect species used in biological control programmes, have been investigated. Various concentrations of the applied herbicides decreased the hatching rate and glycogen levels of the eggs which were still going through embryonic development. It was observed that the glycogen level of Pimpla turionellae eggs that were not exposed to herbicides increased after 4 and 8 h during embryonic development, but decreased in the later stages (24 and 30 h). It was found that the hatching rate and glycogen levels throughout the embryonic development decreased with as the concentrations of 2,4-D and maleic hydrazide applied were increased.     

Host selection by Homalodisca vitripennis: the interplay between feeding, egg maturation, egg load, and oviposition

For some phytophagous insects, egg maturation may be dependent on adult feeding. Accordingly, rates of egg maturation may be dependent on the quality and quantity of available food sources. In turn, oviposition behavior could be affected by diet quality via changes in egg load (number of mature eggs carried by a female). Experiments were conducted to determine whether adult feeding may affect oviposition behavior of the glassy-winged sharpshooter, Homalodisca vitripennis. No-choice tests demonstrated that eggs accumulated in glassy-winged sharpshooter abdomens as time since last oviposition increased largely as a function of feeding plant species. In choice tests, glassy-winged sharpshooter females were observed most frequently on the plant species that imparted the greatest egg maturation rate in no-choice tests. Direct tests of the effects of egg load on glassy-winged sharpshooter oviposition behavior found that females were more likely to deposit eggs as egg load increased. Similarly, acceptance of a low-ranked oviposition plant species by female glassy-winged sharpshooters increased with egg load and time since last oviposition. The results indicate that adult feeding affected glassy-winged sharpshooter egg maturation, plant species varied in quality for providing nutrients for egg maturation, and egg load affected oviposition behavior. Thus, the quantity and quality of available feeding plant species may affect glassy-winged sharpshooter egg maturation rates, which in turn may affect the plant species female glassy-winged sharpshooters select for oviposition.     

Aggregative oviposition varies with density in processionary moths—Implications for insect outbreak propensity

1. In gregarious insects, groups commonly originate from females laying eggs in masses and feeding groups are established as soon as larvae hatch. Some group-living insect species may aggregate beyond the individual parent level, such that offspring from two or more egg masses develop within a common resource.
2. Here we show that aggregative oviposition can vary with population density at oviposition and possibly be an important factor in outbreak dynamics of phytophagous insects.
3. We analysed density data with respect to egg mass aggregation for two species of pine processionary moths, Thaumetopoea pinivora (in Sweden 2005–2019) and T. pityocampa (in Spain 1973–1991). Both species lay their eggs in egg masses and feed in groups. During the study periods, insect population density for both species varied by at least an order of magnitude.
4. The two species showed strikingly similar patterns of egg mass aggregation. Egg masses were overdispersed at high population density, with few trees showing a high load of egg masses.
5. Our data suggest that aggregative oviposition can be important in explaining the previously documented higher propensity for outbreaks in insects laying eggs in clusters, compared with those laying individual eggs.

     

Inhibition of the onset of vitellogenesis by abnormal retention of mature eggs in the stick insect,Carausius morosus

Mature eggs stored in the lateral oviducts are unusual in Carausius morosus. When egg laying is experimentally prevented, ovarian production is inhibited by the accumulation of mature eggs within the lateral oviducts. Such storage affects only the very small terminal oocytes and not the vitellogenic follicles, which attain to term and are ovulated. The fact that unilateral retention of eggs affects only the corresponding ovary strongly suggests that the mature eggs themselves inhibit ovarian production in the stick insect without the involvement of an endocrine organ. It is likely that this mode of action, different from that suggested for the housefly or Rhodnius prolixus, is correlated with the independence of ovarian development from the corpus allatum hormone in the stick insect.     

Centrosome inheritance in the parthenogenetic egg of the collembolan Folsomia candida

Unfertilized eggs commonly lack centrioles, which are usually provided by the male gamete at fertilization, and are unable to assemble functional reproducing centrosomes. However, some insect species lay eggs that develop to adulthood without a contribution from sperm. We report that the oocyte of the parthenogenetic collembolan Folsomia candida is able to self-assemble microtubule-based asters in the absence of pre-existing maternal centrosomes. The asters, which develop near the innermost pole of the meiotic apparatus, interact with the female chromatin to form the first mitotic spindle. The appearance of microtubule-based asters in the cytoplasm of the activated Folsomia oocyte might represent a conserved mechanism for centrosome formation during insect parthenogenesis. We also report that the architecture of the female meiotic apparatus and the structure of the mitotic spindles during the early embryonic divisions are unusual in comparison with that of insects.This work was made possible by grants from PAR (University of Siena) and PRIN to G.C.     

丽蝇蛹集金小蜂的生物学及其滞育调控机制研究进展

丽蝇蛹集金小蜂可以蛹外寄生多种双翅目蝇科害虫,在防治卫生害虫及牲畜害虫方面效果显著。该蜂是实验室内研究寄生蜂的理想模式昆虫,其发育学、行为学、生态学以及遗传学等的研究已有70多年的历史,近年来,随着金小蜂基因组测序的完成,系统的RNAi方法和CRISPR-Cas9技术在丽蝇蛹集金小蜂中的成功应用,该蜂现已成为优良的新型遗传学研究模式昆虫。丽蝇蛹集金小蜂以末龄幼虫进行兼性滞育,其滞育由母代经历的短光照和低温决定,研究该蜂的滞育,不仅有助于揭示昆虫滞育的分子机制,也可帮助解决有益昆虫实际应用中的贮存、运输和适时防治等问题,为提高天敌昆虫的应用效果提供技术支撑。本文总结了近年来丽蝇蛹集金小蜂的研究文献,主要介绍了该蜂的基本生物学特征,概括了现代分子生物学前沿技术在金小蜂研究中的应用情况,并重点讨论了丽蝇蛹集金小蜂滞育的最新研究进展,以期为深入金小蜂的滞育研究和促进该蜂其他研究领域的发展提供参考。     

The influence of parthenogenesis-inducing Wolbachia on the oviposition behaviour and sex-specific developmental requirements of autoparasitoid wasps

Parthenogenesis-inducing (PI) Wolbachia are bacteria that cause incipient male eggs of parasitoid wasps to develop as females. These microbes may invade populations and lead to fixation of parthenogenesis. In this study, the consequences of fixation of PI Wolbachia on evolution of behaviour and development were addressed in three species of whitefly parasitoids in the genus Encarsia. Most sexual Encarsia are obligate autoparasitoids. In these haplodiploid wasps, fertilized, female-producing eggs are laid in whitefly or scale insect nymphs, while unfertilized, male-producing eggs are laid in immature parasitoids within the homopteran integument. The oviposition behaviour and progeny developmental requirements of unmated females of a sexual autoparasitoid species were compared with two species that were fixed for PI Wolbachia. All species were treated with antibiotics, effectively ‘curing’ the asexual species of Wolbachia infection. As expected, unmated females of the sexual species, E. pergandiella, laid few eggs in whiteflies relative to immature parasitoids, and males emerged exclusively from parasitoids. In contrast, females of one of the parthenogenetic species, E. formosa, laid few eggs in parasitoids relative to whiteflies, and male progeny emerged exclusively from whiteflies. Females of the second parthenogenetic species, E. hispida, were intermediate in their oviposition behaviour. Females laid eggs in both wasps and whiteflies. However, all of the males except one emerged from the whiteflies. The results suggest that the association of autoparasitoids and PI Wolbachia may result in dramatic evolutionary change of both host selection behaviour and developmental requirements of the infected wasps.     

Life history,immature morphology and adult behavior ofTelenomus solitus [Hymenoptera: Scelionidae]

The results of laboratory studies on the life history, morphology of immature stages, effects of food and water on adult longevity, host age preference for oviposition, parthenogenesis, and adult behavior ofTelenomus solitus Johnson are presented.Telenomus solitus completed its development in 10.8 days (egg, 0.3 day; 1^st instar, 2.7 days; 2^nd instar, 3.7 days; and pupa, 4.1 days). The adults lived longest when provided with both honey and water. Without food, they survived less than 24 h. Parasitization of host eggs varied significantly with their age, being highest in 42 h old eggs. Fewer than 10% of eggs older than 60 h were parasitized successfully. The periodicity of oviposition was unimodal; it occurred throughout the 24 h period and peaked in the late photophase. Females ofT. solitus are arrhenotokous. The behavioral sequences during courtship, mating, and oviposition are described.      

Plant‐mediated effects of butterfly egg deposition on subsequent caterpillar and pupal development,across different species of wild Brassicaceae

1. Herbivory can change plant quality, which may have consequences for interactions between the inducing herbivore and other insect community members. 2. Studies investigating the effects of plant quality on herbivore performance often have neglected the egg stage, and instead introduced larvae onto the plant. Recently, we reported that herbivore oviposition by Pieris brassicae (Linnaeus) (Large Cabbage White Butterfly) reduced the plant quality of Brassica nigra L. (black mustard) for subsequent herbivores. 3. It remains unclear how persistent and common these plant‐mediated effects of oviposition are. Here, five species of wild Brassicaceae were used (B. nigra L., Brassica oleracea L., Sinapis arvensis L., Moricandia arvensis L., and Moricandia moricandioides Boiss). The response to oviposition by the specialist P. brassicae was determined by following the natural sequence of events: oviposition, egg, larval, and pupal development. All tested plant species are known to interact with P. brassicae in nature. Caterpillar, pupal mass, and development time on plants exposed to butterfly eggs were assessed compared with egg‐free plants. 4. It was shown that the plant‐mediated effects of oviposition are not specific for B. nigra but occur in most of the tested plant species except for M. arvensis. However, the strength of the plant‐mediated effect on caterpillar growth depended on plant species. Thus, across different members of the Brassicaceae family, oviposition can influence plant quality and has negative consequences on P. brassicae growth. Further studies are needed to assess to what extent this trait might be phylogenetically conserved.     

HYBRID BREAKDOWN BETWEEN TWO HAPLODIPLOID SPECIES: THE ROLE OF NUCLEAR AND CYTOPLASMIC GENES

A central question in evolutionary biology concerns the population and genetic processes by which new species arise. Here, the genetic basis of hybrid breakdown between two haplodiploid species, Nasonia vitripennis and N. giraulti is investigated. Hybridization between the two species is normally prevented by microorganisms that cause bidirectional incompatibility. However, after elimination of microorganisms, F₁ hybrids females are readily produced (due to haplodiploidy, males develop from unfertilized eggs and are therefore not hybrids). F₁ hybrid females are viable and fecund, but recombinant (haploid) F₂ male offspring suffer from severe hybrid breakdown (larval and pupal mortality). This is typically interpreted as evidence for the existence of different coadapted gene complexes in the two species, which are broken up by recombination. F₂ recombinant eggs were rescued by fertilization with the complete chromosome complement from either species, supporting the view that hybrid lethality genes tend to be recessive. Negative epistatic interactions occur between nuclear genes of the two species, and between cytoplasmically inherited factors (cytoplasmic genes) of giraulti and nuclear genes of vitripennis. Interactions between nuclear genes and cytoplasmic genes are asymmetric. Experiments clearly demonstrate that the latter incompatibility is not due to maternal-effect genes, but to cytoplasmically inherited elements. Nuclear-mitochondrial interactions are possibly involved.     

A test of the preference–performance hypothesis with stream insects: selective oviposition affects the hatching success of caddisfly eggs

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Reproduction and sexual development in a freshwater gastrotrich

Summary Post-embryonic development of parthenogenic eggs of Lepidodermella squammata was studied by light and electron microscopy in animals of known age and reproductive history. Each bilateral gonad initially contains eight cells. No mitotic proliferation occurs during parthenogenic egg development. Germ cells are tightly clustered, have smooth plasma membranes with no interconnections, and are uninucleate. There is no surrounding ovary or oviduct. At hatching, two cells in each gonad are identifiable as parthenogenic eggs. The enlarged nucleolus of the most mature egg has already attained the morphology that persists throughout vitellogenesis, with intertwined granular and fibrillar threads. Less mature eggs have earlier stages of nucleolar development, and lack indications of meiotic events. Parthenogenic eggs enter vitellogenesis singly, with formation of RER and active Golgi complexes, and the accumulation of lipid, yolk, and various granules. The shell is formed in situ, whereas the spines elongate after egg deposition. Most animals produce four parthenogenic eggs, which undergo immediate development (tachyblastic eggs). Resting (opsiblastic) eggs are rare in isolation culture. Both types of eggs are produced only prior to the formation of sperm and primary oocytes. The absence of synaptonemal complexes, which would indicate synapsis of homologous chromosomes in prophase of meiosis I, implies that parthenogenesis is by apomixis in L. squammata.     

Cytological evidence for self-fertilization in Lasaea subviridis (Galeommatacea: Bivalvia)

Summary

Isolated individuals of Lasaea subviridis simultaneously spawn both eggs and sperm. The sperm attach to the eggs by undergoing an acrosomal reaction and a male pronucleus is incorporated into the egg cytoplasm. Two reduction divisions associated with the production of two polar bodies occur before the first cleavage. These results are consistent with self-fertilization, but not with apomictic or meiotic parthenogenesis. The net fertilization efficiency of Lasaea subviridis natural populations is estimated at 99.93%.     

The effects of carbohydrates upon the survival and reproduction of adult female Pimpla turionellae L. (Hym., Ichneumonidae)

In this study the effects of 23 carbohydrates belonging to various groups upon the survival egg production and egg of female adult Pimpla turionellae L. were investigated. The best results among the carbohydrates tested was obtained with sucrose which was also employed as control. Glucose, maltose, trehalose and melezitose on the other hand showed no significant effect. The egg production was observed to be unaffected by glucose and maltose although it showed significant increase with trehalose and significant decrease with melezitose. Fructose and sorbitol caused a significant decrease in the survival of the insect. Fructose and sorbitol did not have any significant effect upon egg production whereas galactose caused a significant decrease. With the exception of galactose, no carbohydrate caused any significant effect upon egg hatching. Although they did not produce any eggs the female insects survived for 13.17, 15.13, 11.58 and 15.83 days in mannose, melibiose, raffinose and mannitol, respectively. The shortest life span was observed in arabinose followed by α-methyl- D -glucoside, dulcitol, rhamnose, cellobiose, xylose, starch, lactose, sarbose, ribose and glycogen.     

Oviposition choice by the Onopordum capitulum weevil Larinus latus (Coleoptera: Curculionidae) and its effect on the survival of immature stages

Female Larinus latus lay eggs into the capitula of their Onopordum spp. host plants from the onset of capitulum development until the completion of flowering. This tight linkage between insect life-history and plant development optimised larval survival, apart from some initial asynchrony between the readiness of the females to oviposit and the suitability of the very small capitula to physically support eggs. Eggs were laid on bracts and stems of capitula or directly in the florets, when these became available. Both the location of the egg and time of oviposition influenced mortality factors such as egg and larval parasitism, egg desiccation and larval establishment. Overall, survival of eggs laid later in the season into florets was higher, although the adults that emerged were smaller. The change in choice of oviposition site that occurs at the onset of flowering, not only favours survival, but ensures more efficient resource use by the larvae. This occurs because sequential flowering of Onopordum spp. drives a sequential oviposition pattern and spreads the egg load more evenly over the available capitula, reversing a trend earlier in the season to clump eggs that were laid directly on the bracts. Other potential constraints, such as capitulum size, absolute egg density and previous egg-laying do not play a major role in determining the oviposition pattern of L. latus. Larval survival was positively correlated with capitulum size and not strongly influenced by egg density. Competition for resources only appeared to play a role in smaller capitula, and manifested itself in a reduction in the size of emerging adults rather than the death of immatures. In the absence of strong interspecific competition, the oviposition behavior of L. latus has become geared to maximising resource use for larvae (i.e. its own potential competitiveness) rather than minimising interactions with other members of the capitulum endophage guild.     

Predators induce egg retention in prey

To prevent predation on their eggs, prey often avoid patches occupied by predators. As a result, they need to delay oviposition until they reach predator-free patches. Because many species allocate energy to egg production in a continuous fashion, it is not clear what kind of mechanism prey use to delay oviposition. We used females of the phytoseiid mite Neoseiulus cucumeris to study these mechanisms. Females were placed in patches with pollen, a food source they use for egg production, and they were exposed to another phytoseiid mite, Iphiseius degenerans, which is an intraguild predator of N. cucumeris juveniles. We found that the oviposition of N. cucumeris females on patches with the predator was lower than on patches without the predator. Cues left by the intraguild predator were not sufficient to elicit such behaviour. Females of N. cucumeris reduced oviposition when exposed to the predator by retaining the egg inside their body, resulting in a lower developmental rate once these eggs were laid. Hence, females are capable of retaining eggs, but the development of these eggs continues inside the mother’s body. In this way, females gain some time to search for less risky oviposition sites.     

Counterstrategy to Egg Dumping in a Coreid Bug: Recipient Individuals Discard Eggs from Their Backs

The golden egg bug, Phyllomorpha laciniata Vill. (Heteroptera: Coreidae), is the only terrestrial insect in which females oviposit on the backs of female and male conspecifics. Eggs do not survive unless carried by a bug. Herein, I report laboratory observations that egg-carrying individuals actively brush their backs against the host plant seemingly in an effort to rub off eggs. Egg scraping is more common among individuals carrying many eggs than among those carrying only a few eggs. The most recently received eggs were rubbed off first. Females did not avoid laying eggs on the backs of egg-loaded individuals, nor did bugs carrying several eggs resist oviposition attempts more often than unloaded ones. Some males were likely to have fertilized the eggs they scraped off their backs. Laboratory results of active egg removal correspond with egg loss in the field, suggesting that egg scraping may explain egg loss in nature. The data indicate a cost of egg carrying to an individual and an evolutionary arms race between oviposition and discarding behavior.     

Reciprocal inheritance of centrosomes in the parthenogenetic hymenopteran Nasonia vitripennis

Background: In the majority of animals, the centrosome-the microtubule-organizing center of the cell-is assembled from components of both the sperm and the egg. How the males of the insect order Hymenoptera acquire centrosomes is a mystery, as they originate from virgin birth.Results: To address this issue, we observed centrosome, spindle and nuclear behavior in real time during early development in the parthenogenetic hymenopteran Nasonia vitripennis. Female meiosis was identical in unfertilized eggs. Centrosomes were assembled before the first mitotic division but were inherited differently in unfertilized and fertilized eggs. In both, large numbers of asters appeared at the cortex of the egg after completion of meiosis. In unfertilized eggs, the asters migrated inwards and two of them became stably associated with the female pronucleus and the remaining cytoplasmic asters rapidly disappeared. In fertilized eggs, the Nasonia sperm brought in paternally derived centrosomes, similar to Drosophila melanogaster. At pronuclear fusion, the diploid zygotic nucleus was associated only with paternally derived centrosomes. None of the cytoplasmic asters associated with the zygotic nucleus and, as in unfertilized eggs, they rapidly degenerated.Conclusions: Selection and migration of the female pronucleus is independent of the sperm and its aster. Unfertilized male eggs inherit maternal centrosomes whereas fertilized female eggs inherit paternal centrosomes. This is the first system described in which centrosomes are reciprocally inherited. The results suggest the existence of a previously undescribed mechanism for regulating centrosome number in the early embryo.     

Oviposition Deterrence Is Likely an Effect,Not a Mechanism,in the Leaf Beetle <Emphasis Type="Italic">Chrysophtharta bimaculata</Emphasis> (Olivier) (Coleoptera: Chrysomelidae)

Oviposition deterrence is common in many insects as an evolutionary mechanism to reduce subsequent larval competition. We investigated a suspected case of oviposition deterrence by the paropsine chrysomelid, Chrysophtharta bimaculata. In paired choice tests, gravid females were found to prefer ovipositing on host leaves without conspecific eggs, confirming the presence of an apparent oviposition deterrence mechanism. Washing egg batches in water, hexane, or ethanol did not change this preference, suggesting that a soluble marking pheromone was not involved. Furthermore, it is unlikely that a plant-derived oviposition deterring substance is produced as beetles showed no significant oviposition preference between leaves which had been oviposited upon, but then had the eggs removed, and those that had never been oviposited upon. In trials using artificial leaves and mimic egg batches, “leaves” with “egg batches” placed near the tip of the leaf (the preferred site of oviposition in this species) were significantly less likely to be laid upon than artificial leaves where mimic eggs were placed away from the tip. In combination, the results strongly infer that oviposition deterrence in C. bimaculata is due to the mechanical blocking of the oviposition site by the first laid egg-batch, rather than a specific oviposition deterring cue. The apparent oviposition deterrence in this insect may well be an outcome or evolutionary effect of oviposition-site selection, rather than a clear adaptive mechanism to decrease larval competition.