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Insect cuticle exhibits a wide array of micro- and nano-structures in terms of size, form, and function. However, the investigation of cellular mechanisms of morphogenesis has centered around a small number of structure types and organisms. The recent expansion of the taxa studied, and subsequent discoveries prompt us to revisit well-known models, like the one for bristle morphogenesis. In addition, common themes are emerging in the morphogenesis of cuticular structures, such as the polyploidy of precursor cells, the role of pigments and cuticular proteins in controlling chitin deposition in space and time, and the role of the apical extracellular matrix in defining the shape of the developing structure. Understanding how these structures are synthesized in biological systems holds promise for bioinspired design., Competing Interests: Declaration of Competing Interest The author declares no competing or financial interests., (Copyright © 2024 Elsevier Inc. All rights reserved.)

More than 50% of all animal species are insects that undergo complete metamorphosis. The key innovation of these holometabolous insects is a pupal stage between the larva and adult when most structures are completely rebuilt. Why this extreme lifestyle evolved is unclear. Here, we test the hypothesis that a trade-off between growth and differentiation explains the evolution of this novelty. Using a comparative approach, we find that holometabolous insects grow much faster than hemimetabolous insects. Using a theoretical model, we then show how holometaboly evolves under a growth-differentiation trade-off and identify conditions under which such temporal decoupling of growth and differentiation is favored. Our work supports the notion that the holometabolous life history evolved to remove developmental constraints on fast growth, primarily under high mortality., Competing Interests: Competing interests statement:The authors declare no competing interest.

Metabolic theories in ecology interpret ecological patterns at different levels through the lens of metabolism, typically applying allometric power scaling laws to describe rates of energy use. This requires a sound theory for metabolism at the individual level. Commonly used mechanistic growth models lack some potentially important aspects and fail to accurately capture a growth pattern often observed in insects. Recently, a new model (MGM-the Maintenance-Growth Model) was developed for ontogenetic and post-mature growth, based on an energy balance that expresses growth as the net result of assimilation and metabolic costs for maintenance and feeding. The most important contributions of MGM are: 1) the division of maintenance costs into a non-negotiable and a negotiable part, potentially resulting in maintenance costs that increase faster than linearly with mass and are regulated in response to food restriction; 2) differentiated energy allocation strategies between sexes and 3) explicit description of costs for finding and processing food. MGM may also account for effects of body composition and type of growth at the cellular level. The model was here calibrated and evaluated using empirical data from an experiment on house crickets growing under ad libitum conditions. The procedure involved parameter estimations from the literature and collected data, using statistical models to account for individual variation in parameter values. It was found that ingestion rate cannot be generally described by a simple allometry, here requiring a more complex description after maturity. Neither could feeding costs be related to ingestion rate in a simplistic manner. By the unusual feature of maintenance costs increasing faster than linearly with body mass, MGM could well capture the differentiated growth patterns of male and female crickets. Some other mechanistic growth models have been able to provide good predictions of insect growth during early ontogeny, but MGM may accurately describe the trajectory until terminated growth., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Mauritsson, Jonsson. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Molt-based transitions in form are a central feature of insect life that have enabled adaptation to diverse and changing environments. The endocrine regulation of these transitions is well established, but an understanding of their genetic regulation has only recently emerged from insect models. The pupal and adult stages of metamorphosing insects are determined by the stage specifying transcription factors broad-complex (br) and Ecdysone inducible protein 93 (E93), respectively. A probable larval determinant, chronologically inappropriate metamorphosis (chinmo), has just recently been characterized. Expression of these three transcription factors in the metamorphosing insects is regulated by juvenile hormone with ecdysteroid hormones, and by mutual repression between the stage-specific transcription factors. This review explores the hypothesis that variations in the onset, duration, and tissue-specific expression of chinmo, br, and E93 underlie other polyphenisms that have arisen throughout insects, including the castes of social insects, aquatic stages of mayflies, and the neoteny of endoparasites. The mechanisms that constrain how chinmo, br, and E93 expression may vary will also constrain the ways that insect life history may evolve. I find that four types of expression changes are associated with novel insect forms: (1) heterochronic shift in the turnover of expression, (2) expansion or contraction of expression, (3) tissue-specific expression, and (4) redeployment of stage-specific expression. While there is more to be learned about chinmo, br, and E93 function in diverse insect taxa, the studies outlined here show that insect stages are modular units in developmental time and a substrate for evolutionary forces to act upon., (© 2024 The Author(s). Evolution & Development published by Wiley Periodicals LLC.)

Eusocial insects belong to distinct queen and worker castes, which, in turn, can be divided into several morphologically specialized castes of workers. Caste determination typically occurs by differential nutrition of developing larvae. We present a model for the coevolution of larval signalling and worker task allocation-both modelled by flexible smooth reaction norms-to investigate the evolution of caste determination mechanisms and worker polymorphism. In our model, larvae evolve to signal their nutritional state to workers. The workers evolve to allocate time to foraging for resources versus feeding the brood, conditional on the larval signals and their body size. Worker polymorphism evolves under accelerating foraging returns of increasing body size, which causes selection to favour large foraging and small nursing workers. Worker castes emerge because larvae evolve to amplify their signals after obtaining some food, which causes them to receive more food, while the other larvae remain unfed. This leads to symmetry-breaking among the larvae, which are either well-nourished or malnourished, thus emerging as small or large workers. Our model demonstrates the evolution of nutrition-dependent caste determination and worker polymorphism by a self-reinforcement mechanism that evolves from the interplay of larval signalling and worker response to the signals.

Insects can display a vast repertoire of complex and adaptive behaviors crucial for survival and reproduction. Yet, how the neural circuits underlying insect behaviors are assembled throughout development and remodeled during evolution remains largely obscure. The advent of single-cell transcriptomics has opened new paths to illuminate these historically intractable questions. Insect behavior is governed by its brain, whose functional complexity is realized through operations across multiple levels, from the molecular and cellular to the circuit and organ. Single-cell transcriptomics enables dissecting brain functions across all these levels and allows tracking regulatory dynamics throughout development and under perturbation. In this review, we mainly focus on the achievements of single-cell transcriptomics in dissecting the molecular and cellular architectures of nervous systems in representative insects, then discuss its applications in tracking the developmental trajectory and functional evolution of insect brains., Competing Interests: Declaration of Competing Interest We declare that we have no conflict of interest., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Insect growth regulators (IGRs) have been developed as effective control measures against harmful insect pests to disrupt their normal development. This study is to propose a mathematical model to evaluate the cost-effectiveness of IGRs for pest management. The key features of the model include the temperature-dependent growth of insects and realistic impulsive IGRs releasing strategies. The impulsive releases are carefully modeled by counting the number of implements during an insect's temperature-dependent development duration, which introduces a surviving probability determined by a product of terms corresponding to each release. Dynamical behavior of the model is illustrated through dynamical system analysis and a threshold-type result is established in terms of the net reproduction number. Further numerical simulations are performed to quantitatively evaluate the effectiveness of IGRs to control populations of harmful insect pests. It is interesting to observe that the time-changing environment plays an important role in determining an optimal pest control scheme with appropriate release frequencies and time instants., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

For centuries, it has been understood that the final size of adult holometabolous insects is determined by the end of the larval stage, and that once they transform to adults, holometabolous insects do not grow. Despite this, no previous study has directly tested these "old truths" across holometabolous insects. Here, we demonstrate that final adult size is set at the end of the last larval stage in species representing each of the four orders of holometabolous insects: the fruit fly Drosophila melanogaster (Diptera), the tobacco hornworm Manduca sexta (Lepidoptera), the dung beetle Onthophagus taurus (Coleoptera), and the Florida carpenter ant Camponotus floridanus (Hymenoptera). Furthermore, in both D. melanogaster and C. floridanus, we show that the size of adult individuals fluctuates but does not significantly change. Therefore, our study finally confirms these two basic assumptions in the biology of insects, which have for centuries served as the foundation for studies of insect growth, size, and allometry., (© 2022 Wiley Periodicals LLC.)

Ecdysone receptor (EcR) and chitinase play a critical role in the molting stage of insect pests. Each of them is considered a promising target for the development of novel insect growth regulators (IGRs). In the present paper, a total of 24 (23 novel) hexacyclic pyrazolamide derivatives were designed and synthesized by reducing the heptacycle and inserting small flexible linkers on the basis of the previously discovered dual-target compound D-27 acting simultaneously on EcR and Ostrinia furnacalis chitinase ( Of ChtI). Their insecticidal activities against Plutella xylostella , Spodoptera frugiperda , and Ostrinia furnacalis larvae were evaluated. The results revealed that the insecticidal activity was not significantly enhanced when the heptacycle on the pyrazole ring was reduced to a hexacycle. However, the insertion of an additional methylene spacer between the substituted phenyl ring and the amide bond can improve the insecticidal activity. Among the derivatives, the most potent compound, 6j , exhibited promising insecticidal activities against P. xylostella and S. frugiperda . Further protein binding assays and molecular docking indicated that 6j could target both EcR and Of ChtI, and is a potential lead compound for IGRs. The present work provides valuable clues for the development of new dual-target IGRs.

Metazoans function as individual organisms but also as "colonies" of cells whose single-celled ancestors lived and reproduced independently. Insights from evolutionary biology about multicellular group formation help us understand the behavior of cells: why they cooperate, and why cooperation sometimes breaks down. Current explanations for multicellularity focus on two aspects of development which promote cooperation and limit conflict among cells: a single-cell bottleneck, which creates organisms composed of clones, and a separation of somatic and germ cell lineages, which reduces the selective advantage of cheating. However, many obligately multicellular organisms thrive with neither, creating the potential for within-organism conflict. Here, we argue that the prevalence of such organisms throughout the Metazoa requires us to refine our preconceptions of conflict-free multicellularity. Evolutionary theory must incorporate developmental mechanisms across a broad range of organisms-such as unusual reproductive strategies, totipotency, and cell competition-while developmental biology must incorporate evolutionary principles. To facilitate this cross-disciplinary approach, we provide a conceptual overview from evolutionary biology for developmental biologists, using analogous examples in the well-studied social insects.

Bacillus cereus is a spore forming bacteria recognized among the leading agents responsible for foodborne outbreaks in Europe. B. cereus is also gaining notoriety as an opportunistic human pathogen inducing local and systemic infections. The real incidence of such infection is likely underestimated and information on genetic and phenotypic characteristics of the incriminated strains is generally scarce. We have recently analyzed a large strain collection of varying pathogenic potential. Screening for biomarkers to differentiate among clinical and non-clinical strains, a gene encoding an alcohol dehydrogenase-like protein was identified among the leading candidates. This family of proteins has been demonstrated to be involved in the virulence of several bacterial species. The relevant gene was knocked out to elucidate its function with regards to resistance to host innate immune response, both in vitro and in vivo. Our results demonstrate that the adhB gene plays a significant role in resistance to nitric oxide and oxidative stress in vitro, as well as its pathogenic ability with regards to in vivo toxicity. These properties may explain the pathogenic potential of strains carrying this newly identified virulence factor., Competing Interests: The authors have declared that no competing interests exist.

DNA methylation is a process in which methyl (CH3) groups are added to the DNA molecule. The DNA segment does not change in the sequence, but DNA methylation could alter the action of DNA. Different enzymes like DNA methyltransferases (DNMTs) take part in methylation of cytosine/adenine nucleosides in DNA. In prokaryotes, DNA methylation is performed to prevent the attack of phage and also plays a role in the chromosome replication and repair. In fungi, DNA methylation is studied to see the transcriptional changes, as in insects, the DNA methylation is not that well-known, it plays a different role like other organisms. In mammals, the DNA methylation is related to different types of cancers and plays the most important role in the placental development and abnormal DNA methylation connected with diseases like cancer, autoimmune diseases, and rheumatoid arthritis.

The development rates of arthropods are temperature-dependent. Studies aiming to predict the dynamics of arachnid, crustacean, and insect populations in nature often require the derivation of development functions representing this phenomenon. A previous study (Quinn, B.K., 2017, J. Therm. Biol. 63, 65-77) identified 33 development functions commonly used in past studies on temperature-dependent development of arthropods, and illustrated that: (1) most of 99 past studies only applied one or few (2-5) development functions to their data without considering others; and (2) most of a subset of 79 studies' data were not fit with the actual best function for them, resulting in sometimes substantial differences in model performance and predictive ability. However, that study did not test the class of development functions based on theoretical enzyme thermodynamics, including the Sharpe-Schoolfield-Ikemoto (SSI) function. Herein, the meta-analyses done in that previous study were redone, after fitting all 79 reanalyzed datasets with the SSI function. Estimates of the intrinsic optimum temperature (T Φ ) for development of each tested species were also derived using the SSI function and compared among taxa. Including the SSI function in analyses did not change the conclusions of the previous study concerning development function usage, choice, and consequences. Notably, the SSI function performed as well as or relatively better than other functions of comparable or lower complexity in terms of R 2 , AIC C -based rankings, ΔAIC C values, and prediction errors, which may recommend its more widespread use in future studies. Overall differences in T Φ were found among arthropod subphyla, as well as between most species pairs. Most T Φ estimates produced herein were novel, and could be used to make inferences about or comparisons among arthropod taxa in future studies., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Carrion is a dynamic and nutrient-rich resource that attracts numerous insect species that undergo succession due to the rapid change in the carrion resource. Despite this process being well-understood, few studies have examined resource change as a driver of carrion insect succession, and instead have focused on the effects of time per se, or on coarse, qualitative measures such as decay stage. Here we report on three field succession experiments using pig carcasses and human cadavers encompassing two winters and one summer. We quantified the effects of resource change (measured as total body score, TBS), carrion type, initial carrion mass, ambient temperature, and season on insect species richness and community composition. We found that all variables had an effect on different taxonomic or trophic components of the insect community composition, with the exception of initial carrion mass which had no effect. We found significant positive effects of TBS on beetle species richness and composition, while fly species richness was not significantly affected by TBS, but was by ambient temperature. TBS had a significant positive effect on all trophic groups, while ambient temperature also had a significant positive effect on the necrophages and predator/parasitoids. Our study indicates that resource change, as indicated by TBS, is an important driver of carrion insect species turnover and succession on carrion, and that TBS can provide information about insect ecological patterns on carrion that other temporal measures of change cannot., (© The Author(s) 2021. Published by Oxford University Press on behalf of Entomological Society of America.All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Most animal species consist of two distinct sexes. At the morphological, physiological, and behavioral levels the differences between males and females are numerous and dramatic, yet at the genomic level they are often slight or absent. This disconnect is overcome because simple genetic differences or environmental signals are able to direct the sex-specific expression of a shared genome. A canonical picture of how this process works in insects emerged from decades of work on Drosophila. But recent years have seen an explosion of molecular-genetic and developmental work on a broad range of insects. Drawing these studies together, we describe the evolution of sexual dimorphism from a comparative perspective and argue that insect sex determination and differentiation systems are composites of rapidly evolving and highly conserved elements., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Acquisition of novel structures often has a profound impact on the adaptation of organisms. The wing of insects is one such example, facilitating their massive success and enabling them to become the dominant clade on this planet. However, its evolutionary origin as well as the mechanisms underpinning its evolution remain elusive. Studies in crustaceans, a wingless sister group of insects, have played a pivotal role in the wing origin debate. Three recent investigations into the genes related to insect wings and legs in crustaceans provided intriguing insights into how and where insect wings evolved. Interestingly, each study proposes a distinct mechanism as a key process underlying insect wing evolution. Here, I discuss what we can learn about the evolution of insect wings and morphological novelty in general by synthesizing the outcomes of these studies., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Moulting is a cornerstone of arthropods development. It can be determined by numerous factors such as body mass, temperature, and immunity. However, the effects of these factors can be dependent on each other, so that it is often difficult to predict whether and how they shape moulting, and whether their effects are additive or interactive. In this study, we addressed these questions by testing the effects of body mass, ambient temperature, fungal infection and their interaction on intermoult duration and survival in starved juveniles of the European earwig Forficula auricularia. We recorded the date of moult and death of a total of 207 earwig juveniles that were weighed, exposed to different doses of the entomopathogenic fungus Metarizium brunneum and then maintained at either 20 °C or 24 °C. Our results first reveal that juveniles moulted earlier when they were heavy compared to light on the day of exposure, as well as earlier when maintained at 24 °C compared to 20 °C. By contrast, pathogen exposure did not affect the moulting date. We also found that nymphs died faster when they were light compared to heavy on the day of exposure, when they were exposed to high (10 6 and 10 7 spores/ml) compared to low (10 4 , 10 5 and 0 spores/ml) pathogen concentrations, and when they were maintained at 24 °C compared to 20 °C. We detected no sign of interaction between temperature, fungal infection and body mass on both moulting and survival. Overall, these findings shed light on the limited importance of infection on moulting in starved juveniles, and reveal that weight, temperature, and infection have additive effects on their survival. More generally, this study emphasizes that the three tested factors do not necessarily interact to shape key physiological processes in an insect., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Nucleic acids help clarify variation in species richness of insects having different metamorphosis modes, a biological conundrum. Here we analyse nucleic acid contents of 639 specimens of aquatic insects collected from four high mountain streams of Sierra Nevada in southern Spain to test whether the allocation to RNA or DNA content differs during ontogeny between juvenile insects undergoing direct (hemimetabolous) or indirect (holometabolous) metamorphosis. The results show that RNA content as a function of body mass was negatively correlated to insect body length in four out of six and three out of six of the holometabolan and hemimetabolan taxa, respectively. Although no significant differences in RNA content were found between holometabolans and hemimetabolans, the significant interaction between body length and metamorphosis mode for RNA and RNA:DNA indicates a strong ontogenetic component to RNA allocation. In addition, our finding of lower DNA content in holometabolans relative to hemimetabolans agree with the analysis of empirical genome data in aquatic and terrestrial insects, and extend to this class of arthropods the "growth rate-genome size-nutrient limitation" hypothesis that differences in allocation between RNA and DNA may reflect fundamental evolutionary trade-off of life-history strategies associated with high growth rates (and RNA content) in holometabolans at the expense of diminished genome sizes.

A major current challenge is to increase the food production while preserving natural resources. Agricultural practices that enhance the productivity and progressively improve the soil quality are relevant to face this challenge. Trichoderma species are widely used in agriculture to stimulate the plant growth and to control different pathogens affecting crops, representing useful tools for sustainable food production. This mini-review summarizes applications of Trichoderma strains in agriculture to control fungal pathogens, nematodes and insects, the involved biocontrol mechanisms, efficacy and inoculation forms in greenhouse, field and post-harvest conditions. Aspects of Trichoderma handling that influence on biocontrol efficacy such as preventive treatments, frequency of applications and delivery methods are discussed. Strategies useful to improve the antagonistic performance such as the use of native strains, protoplast fusion, formulation, growth on pathogen cell wall medium and combination with other antagonists in integrated treatments are discussed. This mini-review provides practical knowledge to design safe and optimal biocontrol strategies based on Trichoderma and pose challenges to expand its antagonist performance.

There are two major categories of observation data in studying time-dependent processes: one is the time-series data, and the other is the perhaps lesser-recognized but similarly prevalent time-to-event data (also known as survival or failure time). Examples in entomology include molting times and death times of insects, waiting times of predators before the next attack or the hiding times of preys. A particular challenge in analyzing time-to-event data is the observation censoring, or the incomplete observation of survival times, dealing which is a unique advantage of survival analysis statistics. Even with a perfectly designed experiment being conducted perfectly, such 'naturally' censoring may still be unavoidable due to the natural processes, including the premature death in the observation of insect development, the variability in instarship, or simply the continuous nature of time process and the discrete nature of sampling intervals. Here we propose to apply the classic Cox proportional hazards model for modeling both insect development and survival rates (probabilities) with a unified survival analysis approach. We demonstrated the advantages of the proposed approach with the development and survival datasets of 1800 Russian wheat aphids from their births to deaths, observed under 25 laboratory treatments of temperatures and plant growth stages.

Glycosyltransferases (GTs) catalyse the reaction of glyco-conjugation of various biomolecules by transferring the saccharide moieties from an activated nucleotide sugar to nucleophilic glycosyl acceptor. In insects, GTs show diverse temporal and site-specific expression patterns and thus play significant roles in forming the complex biomolecular structures that are necessary for insect survival, growth and development. Several insects exhibit GT-mediated detoxification as a key defence strategy against plant allelochemicals and xenobiotic compounds, as well as a mechanism for pesticide cross-resistance. Also, these enzymes act as crucial effectors and modulators in various developmental processes of insects such as eye development, UV shielding, cuticle formation, epithelial development and other specialized functions. Furthermore, many of the known insect GTs have been shown to play a fundamental role in other physiological processes like body pigmentation, cuticular tanning, chemosensation and stress response. This review provides a detailed overview of the multifaceted functionality of insect GTs and summarizes numerous case studies associated with it., (© 2020 The Royal Entomological Society.)

Metamorphosis undeniably shaped the evolutionary success of winged insects. So far, what we know about the hormonal regulation and molecular mechanisms controlling insect metamorphosis lies on the understanding of complete and incomplete metamorphosis. Rarer types of metamorphosis are overlooked, yet they could provide important insights as they represent deviations in life history strategies that are associated with unique ecological traits. The molecular mechanisms of these atypical metamorphoses are still poorly understood. With the rise of next-generation sequencing, and increasing interest in emerging organismal systems, it is now possible to start exploring the molecular mechanisms underlying atypical metamorphoses in insects. By focusing on neometaboly and paedomorphosis, we discuss how exploring their molecular mechanisms can complete our understanding on the evolution of insects and impact applied research areas. Continued decrease in next-generation sequencing costs and progress in genome editing will help decipher the proximate mechanisms of unusual life history strategies in insects., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Ecdysteroids are a class of steroid hormones that controls molting and metamorphic transitions in Ecdysozoan species including insects, in which ecdysteroid biosynthesis and its regulation have been extensively studied. Insect ecdysteroids are produced from dietary sterols by a series of reduction-oxidation reactions in the prothoracic gland and in Drosophila they are released into the hemolymph via vesicle-mediated secretion at the time of metamorphosis. To initiate precisely controlled ecdysteroid pulses, the prothoracic gland functions as a central node integrating both intrinsic and extrinsic signals to control ecdysteroid biosynthesis and secretion. In this review, we outline recent progress in the characterization of ecdysone biosynthesis and steroid trafficking pathways and the discoveries of novel factors regulating prothoracic gland function., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Proper formation of adult insects requires the integration of spatial and temporal regulatory axes. Whereas spatial information confers identity to each tissue, organ and appendage, temporal information specifies at which stage of development the animal is. Regardless of the type of post-embryonic development, either hemimetabolous or holometabolous, temporal specificity is achieved through interactions between the temporal identity genes Kr-h1, E93 and Br-C, whose sequential expression is controlled by the two major developmental hormones, 20-hydroxyecdysone and Juvenile hormone. Given the intimate regulatory connection between these three factors to specify life stage identity, we dubbed the regulatory axis that comprises these genes as the Metamorphic Gene Network (MGN). In this review, we survey the molecular mechanisms underlying the control by the MGN of stage identity and progression in hemimetabolous and holometabolous insects., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Metamorphosis is fascinating and dramatic stage of postembryonic development in insects [1]. The most prominent metamorphic changes seen in holometabolous insects involve destruction of most larval structures and concomitant generation of adult ones. Such diverse cellular events are orchestrated by ecdysone. The central nervous system (CNS) is also extensively remodeled to process new sensory inputs; to coordinate new types of locomotion; and to perform higher-order decision making [2]. Programmed cell death (PCD) is an integral part of the metamorphic development. It eliminates obsolete larval tissues and extra cells that are generated from the morphogenesis of adult tissues. In the CNS, PCD of selected neurons and glial cells as well as reshaping of persistent larval cells are essential for establishing the adult CNS. In this review, we summarize the ecdysone signaling, and then molecular and cellular events associated with PCD primarily in the metamorphosing CNS of Drosophila melanogaster., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Metamorphosis (Greek for a state of transcending-form or change-in-shape) refers to a dramatic transformation of an animal's body structure that occurs after development of the embryo or larva in many species. The development of a fly (or butterfly) from a crawling larva (or caterpillar) that forms a pupa (or chrysalis) before eclosing as a flying adult is a classic example of metamorphosis that captures the imagination and has been immortalized in children's books. Powerful genetic experiments in the fruit fly Drosophila melanogaster have revealed how genes can instruct the behaviour of individual cells to control patterns of tissue growth, mechanical force, cell-cell adhesion and cell-matrix adhesion drive morphogenetic change in epithelial tissues. Together, the distribution of mass, force and resistance determines cell shape changes, cell-cell rearrangements, and/or the orientation of cell divisions to generate the final form of the tissue. In organising tissue shape, genes harness the power of self-organisation to determine the collective behaviour of molecules and cells, which can often be reproduced in computer simulations of cell polarity and/or tissue mechanics. This review highlights fundamental discoveries in epithelial morphogenesis made by pioneers who were fascinated by metamorphosis, including D'Arcy Thompson, Conrad Waddington, Dianne Fristrom and Antonio Garcia-Bellido., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Modifications to DNA and core histones influence chromatin organization and expression of the genome. DNA methylation plays a significant role in the regulation of multiple biological processes that regulate behavior and caste differentiation in social insects. Histone modifications play significant roles in the regulation of development and reproduction in other insects. Genes coding for acetyltransferases, deacetylases, methyltransferases, and demethylases that modify core histones have been identified in genomes of multiple insects. Studies on the function and mechanisms of action of some of these enzymes uncovered their contribution to post-embryonic development. The results from studies on epigenetic modifiers could help in the identification of inhibitors of epigenetic modifiers that could be developed to control pests and disease vectors., (Copyright © 2020 Elsevier Inc. All rights reserved.)

All organisms are exposed to changes in their environment throughout their life cycle. When confronted with these changes, they adjust their development and physiology to ensure that they can produce the functional structures necessary for survival and reproduction. While some traits are remarkably invariant, or robust, across environmental conditions, others show high degrees of variation, known as plasticity. Generally, developmental processes that establish cell identity are thought to be robust to environmental perturbation, while those relating to body and organ growth show greater degrees of plasticity. However, examples of plastic patterning and robust organ growth demonstrate that this is not a hard-and-fast rule.In this review, we explore how the developmental context and the gene regulatory mechanisms underlying trait formation determine the impacts of the environment on development in insects. Furthermore, we outline future issues that need to be resolved to understand how the structure of signaling networks defines whether a trait displays plasticity or robustness.

While nanoparticles (NPs) can be used as insecticides by themselves, they can also be carriers for insecticidal chemicals. Existing literature suggests that the smaller the NP size, the greater the toxicity and penetration into the insect's body. Nonetheless, there is a lack of literature pertaining to the mode of action within insects. This review article summarizes the currently available entomological studies on the mechanisms of NP-insect interactions. Externally, NPs affect pigmentation and integrity of the cuticle, while internally they induce immune responses and alter gene expression leading to altered protein, lipid, and carbohydrate metabolism along with cellular toxicity that impairs development and reproduction of the insect. Consequently, insects are incapacitated due to the disruption of the nutrient intake, production of reactive oxygen species and altered biochemical activity while some NPs can promote growth and development as well as diminish the effects of nontarget toxicity.

Spray application is considered to be the most common method of insecticide use in apple orchard, while trunk injection has often be used in labor-intensive areas. Here, a comparison of both methods in aphid control efficiency and non-target effects was conducted. We evaluated the effects of thiamethoxam by either spray or injection on apple aphid Aphis citricola, and examined the temporal dynamic of thiamethoxam in leaves by using residue analysis. Results showed that thiamethoxam had a remarkable suppression effect on Aphis citricola, and both application methods had obvious control efficiency with the highest value above 90%. The control effect of spray method on Aphis citricola reached the maximum at 7 days after application, while that of injection method reached the maximum at 14 days after application. Moreover, the control effect of injection after 14 days and that of spray after 7 days were not significant, suggesting the spray method had a higher quick-acting effect than the injection method, and the two methods had a similar persistence effect. The population dynamics of non-target insects (ladybugs, parasitoid wasps and predatory bugs) showed basically the same as that of blank controlled. The control effect evaluation of thiamethoxam on Aphis citricola suggest that injection treatment was more effective in protecting natural enemies than spray treatment, and thiamethoxam didn't interfere with natural enemies to control Aphis citricola with both two application methods., (Copyright © 2020 Elsevier Inc. All rights reserved.)

The recent introduction of the Asian yellow-legged hornet, Vespa velutina, into Europe has raised concern regarding the threat to honeybees and the competition with the European hornet, Vespa crabro . The aim of this study was to investigated essential (Mg, Fe, Zn, Cu) and non-essential (Cd and Pb) elements in these two species. Element concentrations were determined in the whole body and separately in the head, thorax and abdomen using atomic absorption spectrometry (AAS). The changes in essential element concentration and speciation during metamorphosis were also studied using size exclusion chromatography followed by AAS and proteomic analysis. In both species, the essential elements were more concentrated in the abdomen due to the presence of fat bodies. Magnesium, Fe and Zn concentrations were significantly higher in V . crabro than in V. velutina and could have been related to the higher aerobic energy demand of the former species required to sustain foraging flight. Low concentrations of Cd and Pb were indicative of low environmental exposure. The concentration and speciation of essential elements, particularly Fe, varied among the developmental stages, indicating a modification of ligand preferences during metamorphosis. Overall, the results in the present study provide a better understanding of the hornet metal metabolism and a foundation for additional studies.

Zygentoma occupies a key evolutionary position for understanding the evolution of insect metamorphosis but has received little attention in terms of genetic analysis. To develop functional genomic studies in this insect, we evaluated five candidate internal reference genes for quantitative RT-PCR (qPCR) studies from Thermobia domestica , a representative species of Zygentoma, including Actin 5C ( Actin5C ), Elongation factor-1 alpha ( EF1A ), Ribosome protein S26 ( RPS26 ), Ribosome protein L32 ( RPL32 ), and Superoxide dismutase 2 ( SOD2 ), at different developmental stages, in various body parts, and under dsRNA microinjection and starvation stresses, using four algorithms (delta Ct, geNorm, NormFinder and BestKeeper) and a comparative algorithm (RefFinder). Specific suitable reference genes were recommended across specific experimental conditions, and the combination of RPS26 and RPL32 was appropriate for all tested samples. Employing our selected reference gene combination, we investigated the gene expression pattern of Myoglianin ( Myo ), a crucial gene-regulating insect metamorphosis, in ametabolous T. domestica , and demonstrated the efficiency of RNA interference (RNAi) in firebrat nymphs. This study provides a basis for reliable quantitative studies of genes and greatly benefits evolutionary and functional genomics studies in Zygentoma.

The discovery of novel, effective, and botanical pesticides is one of the main strategies for modern plant protection and insect pest control. During the search for novel botanical pesticides from natural sources, the seeds of Sophora tonkinensis were systematically investigated to obtain 11 new matrine-type alkaloids ( 1 - 11 ), including one novel matrine-type alkaloid featuring an unprecedented 5/6/6/6 tetracyclic skeleton ( 1 ), along with 16 known compounds ( 12 - 27 ). Their structures were elucidated by comprehensive spectroscopic data analysis (IR, UV, NMR, and HRESIMS), ECD calculations, and single-crystal X-ray diffraction. The anti-tobacco mosaic virus (TMV) activity and insecticidal activities against Aphis fabae and Tetranychus urticae of the compounds were also respectively screened using the half-leaf method and spray method. Biological tests indicated that compounds 2 , 4 , 6 , and 26 displayed significant anti-TMV biological activities compared with the positive control ningnanmycin. Compounds 7 , 17 , and 26 presented moderate activities against A. fabae with LC 50 values of 38.29, 18.63, and 23.74 mg/L, respectively. Moreover, compounds 13 and 26 exhibited weak activities against T. urticae .

Globally, the citrus industry produces various wastes, which contain a great deal of limonoids. In order for the sustainable development of the citrus industry, and considering the diverse bioactivities of limonoids, a series of ester derivatives were constructed by structural modification of limonin in the B ring. Furthermore, two seven-membered lactone derivatives of limonin and obacunone with a novel skeleton in the B ring were obtained by the Baeyer-Villiger oxidation rearrangement. The steric structures of six key compounds 3a , 3b , 4m , 4n , 6 , and 7 were determined by X-ray crystallography. It demonstrated that the molar ratio of 3a (7α-isomer) and 3b (7β-isomer) depended on the mixed solvents in the reduction system. The anti-tobacco mosaic virus (TMV) activities under three different modes of action for most of the tested compounds were as the following sequence: inactivation effect > protection effect > curative effect. It was noteworthy that compound 4aa displayed the most potent anti-TMV/insect growth inhibitory activities, which indicated that the introduction of the phenylacryloyloxy group at the C-7β position of limonin could significantly improve its agricultural biological activities. This study will pave the way for future value-added application of citrus industrial wastes and provide strong evidence for the discovery of sustainable biopesticides based on limonoids.

Over the course of the last few years, the consumption of insects, known as entomophagy, has sparked increasing interest amongst scientists and environmentalists as a potential solution to the inevitable global food security and sustainability issues humans will be facing in the coming years. Despite the fact that insects have been an integral part of over 2 billion people's diet worldwide, the concept of eating insects is still new to Western culture. As a result, there are many unknowns regarding insects as a food source, and this has led to a number of studies and investigations being done in recent years to create more knowledge and awareness around this new concept in the food industry. This review discusses some of the key topics and new developments published over recent years, such as the nutritional benefits, food safety concerns, functional properties, potential product concepts and the current ideas and attitudes towards insects as a food source in Western culture. © 2017 Society of Chemical Industry., (© 2017 Society of Chemical Industry.)

Arthropods operate in an outrageous diversity of environments. From the deep sea to dense tropical forests, to wide open arctic tundra, they have colonized almost every possible habitat. Within these environments, the presence of light is nearly ubiquitous, varying in intensity, wavelength, and polarization. Light provides critical information about the environment, such as time of day or where food sources may be located. Animals take advantage of this prevalent and informative cue to make behavioral choices. However, the types of choices animals face depend greatly on their environments and needs at any given time. In particular, animals that undergo metamorphosis, with arthropods being the prime example, experience dramatic changes in both behavior and ecology, which in turn may require altering the structure and function of sensory systems such as vision. Amphibiotic organisms maintain aquatic lifestyles as juveniles before transitioning to terrestrial lifestyles as adults. However, light behaves differently in water than in air, resulting in distinct aquatic and terrestrial optical environments. Visual changes in response to these optical differences can occur on multiple levels, from corneal structure down to neural organization. In this review, we summarize examples of alterations in the visual systems of amphibiotic larval and adult insects and malacostracan crustaceans, specifically those attributed to environmental differences between metamorphic phases., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Insects have distinct life stages that can differ in their responses to environmental factors. We discuss empirical evidence and theoretical models for ontogenetic variation in thermal sensitivity and performance curves (TPCs). Data on lower thermal limits for development (T 0 ) demonstrate variation between stages within a species that is of comparable magnitude to variation among species; we illustrate the consequences of such ontogenetic variation for developmental responses to changing temperature. Ontogenetic variation in optimal temperatures and upper thermal limits has been reported in some systems, but current data are too limited to identify general patterns. The shapes of TPCs for different fitness components such as juvenile survival, adult fecundity, and generation time differ in characteristic ways, with important consequences for understanding fitness in varying thermal environments. We highlight a theoretical framework for incorporating ontogenetic variation into process-based models of population responses to seasonal variation and climate change., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Natural products are a source of many novel compounds with biological activity for the discovery of new pesticides and pharmaceuticals. Quinoxaline is a fused N -heterocycle in many natural products and synthetic compounds, and seven novel quinoxaline derivatives were designed and synthesized via three steps. Pesticidal activities of title quinoxaline derivatives were bioassayed. Most of these compounds had herbicidal, fungicidal, and insecticidal activities. The compounds 2-(6-methoxy-2-oxo-3-phenylquinoxalin-1(2 H )-yl)acetonitrile ( 3f ) and 1-allyl-6-methoxy-3-phenylquinoxalin-2(1 H )-one ( 3g ) were the most active herbicides and fungicides. Mode-of-action studies indicated that 3f is a protoprophyrinogen oxidase-inhibiting herbicide. Compound 3f also possessed broad-spectrum fungicidal activity against the plant pathogen Colletotrichum species. Some of these compounds also had insecticidal activity. Molecular docking and DFT analysis can potentially be used to design more active compounds.

A diverse set of pathogens have evolved extended phenotypes that manipulate the moribund behavior of their various insect hosts. By elevating host positioning at death, a phenomenon called "summit disease", these pathogens have been shown to have higher fitness. Though a few summit disease systems have been intensively characterized, in particular the Ophiocordyceps-ant system, summit diseases lack an overarching theory for the underlying mechanisms of this complex behavioral manipulation. In this article, we combine the gamut of summiting systems into a cohesive framework: we propose two types of summit disease (juvenile and adult), which both exploit natural insect behaviors during periods of quiescence. We place this framework in the context of available literature and propose investigations that follow from this comprehensive understanding of summit disease in insects., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Background: Metamorphosis remains one of the most complicated and poorly understood processes in insects. This is particularly so for the very dynamic transformations that take place within the pupal sheath of holometabolous insects. Only few studies address these transformations especially with regard to cranial structures of those holometabolous species where the larval and adult forms have a similar diet. It thus remains unclear to what extent the internal structures undergo histolysis and rebuilding. Here, the development of the brain and skeleto-muscular system of the head of Chrysopa pallens (Rambur, 1838) is studied. This species is a predator of aphids in the larval and adult stage., Results: We used micro-computed-tomography (μ-CT) to study the transformations of the larval, prepupal and pupal head within the cocoon. We first assessed the morphological differences and similarities between the stages. We then determined the point in time when the compound eyes appear and describe the re-orientation of the head capsule which transforms the prognathous larva into a hypognathous adult. The internal head muscles are distinctly more slender in larvae than adults. In addition, the adults have a significantly larger brain which is likely needed for the processing of the signals obtained by the adults vastly expanded sensory organs that are presumably needed for dispersal and mating. Our study shows that the histolysis and modification of the inner muscles and skeletal elements take place within the prepupa. The central nervous system persists throughout metamorphosis but its morphology changes significantly., Conclusion: Our study reveals that not only the inner structures, but also the outer morphology continues to change after the final larval moult. The adult cuticle and internal structures form gradually within the cocoon. The histolysis and rebuilding begin with the skeletal elements and is followed by changes in the central nervous system before it concludes with modifications of the musculature. This order of events is likely ancestral for Holometabola because it is also known from Hymenoptera, Diptera, Mecoptera, and Coleoptera.

The reproductive success of many insects is considered to be limited by two main factors: the availability of mature eggs to lay (termed egg limitation) and the time to locate suitable hosts (termed time limitation). High host density in the environment is likely to enhance oviposition opportunities, thereby selecting for higher investment in egg supply. In contrast, a shortage of food (e.g. sugar sources) is likely to increase the risk of time limitation, thereby selecting for higher allocation to initial energy reserves. To our knowledge, the combined effect of host and food availability on these optimal life-history allocations has never been investigated. We thus modelled their simultaneous effects on a three-dimensional trade-off between initial investment in energy reserves, egg number and egg size, while focusing on insect parasitoids. The model was based on Monte Carlo simulations coupled with genetic algorithms, in order to identify the optimal life-history traits of a single simulated parasitoid female in an environment in which both hosts and food are present in varying densities. Our results reproduced the simple predictions described above. However, some novel predictions were also obtained, especially when specific interactions between the different factors were examined and their effects on the three-dimensional life-history surface were considered. The work sheds light on long-lasting debates regarding the relative importance of time versus egg limitation in determining insect life-history traits and highlights the complexity of life-history evolution, where several environmental factors act simultaneously on multiple traits., (© 2020 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2020 European Society For Evolutionary Biology.)

Metamorphic transformation from larvae to adults along with the high fecundity is key to insect success. Insect metamorphosis and reproduction are governed by two critical endocrines, juvenile hormone (JH), and 20-hydroxyecdysone (20E). Recent studies have established a crucial role of microRNA (miRNA) in insect metamorphosis and oogenesis. While miRNAs target genes involved in JH and 20E-signaling pathways, these two hormones reciprocally regulate miRNA expression, forming regulatory loops of miRNA with JH and 20E-signaling cascades. Insect metamorphosis and oogenesis rely on the coordination of hormones, cognate genes, and miRNAs for precise regulation. In addition, the alternative splicing of genes in JH and 20E-signaling pathways has distinct functions in insect metamorphosis and oogenesis. We, therefore, focus in this review on recent advances in post-transcriptional regulation, with the emphasis on the regulatory role of miRNA and alternative splicing, in insect metamorphosis and oogenesis. We will highlight important new findings of miRNA interactions with hormonal signaling and alternative splicing of JH receptor heterodimer gene Taiman.

In insects, 20-hydroxyecdysone (20E) limits the growth period by triggering developmental transitions; 20E also modulates the growth rate by antagonizing insulin/insulin-like growth factor signaling (IIS). Previous work has shown that 20E cross-talks with IIS, but the underlying molecular mechanisms are not fully understood. Here we found that, in both the silkworm Bombyx mori and the fruit fly Drosophila melanogaster , 20E antagonized IIS through the AMP-activated protein kinase (AMPK)-protein phosphatase 2A (PP2A) axis in the fat body and suppressed the growth rate. During Bombyx larval molt or Drosophila pupariation, high levels of 20E activate AMPK, a molecular sensor that maintains energy homeostasis in the insect fat body. In turn, AMPK activates PP2A, which further dephosphorylates insulin receptor and protein kinase B (AKT), thus inhibiting IIS. Activation of the AMPK-PP2A axis and inhibition of IIS in the Drosophila fat body reduced food consumption, resulting in the restriction of growth rate and body weight. Overall, our study revealed an important mechanism by which 20E antagonizes IIS in the insect fat body to restrict the larval growth rate, thereby expanding our understanding of the comprehensive regulatory mechanisms of final body size in animals., Competing Interests: The authors declare no competing interest.

Insect molting hormone (ecdysteroids) and juvenile hormone regulate molting and metamorphic events in a variety of insect species. Mealybugs undergo sexually dimorphic metamorphosis: males develop into winged adults through non-feeding, pupa-like stages called prepupa and pupa, while females emerge as neotenic wingless adults. We previously demonstrated, in the Japanese mealybug Planococcus kraunhiae (Kuwana), that the juvenile hormone titer is higher in males than in females at the end of the juvenile stage, which suggests that juvenile hormone may regulate male-specific adult morphogenesis. Here, we examined the involvement of ecdysteroids in sexually dimorphic metamorphosis. To estimate ecdysteroid titers, quantitative RT-PCR analyses of four Halloween genes encoding for cytochrome P450 monooxygenases in ecdysteroid biosynthesis, i.e., spook, disembodied, shadow and shade, were performed. Overall, their expression levels peaked before each nymphal molt. Transcript levels of spook, disembodied and shadow, genes that catalyze the steps in ecdysteroid biosynthesis in the prothoracic gland, were higher in males from the middle of the second nymphal instar to adult emergence. In contrast, the expression of shade, which was reported to be involved in the conversion of ecdysone into 20-hydroxyecdysone in peripheral tissues, was similar between males and females. These results suggest that ecdysteroid biosynthesis in the prothoracic gland is more active in males than in females, although the final conversion into 20-hydroxyecdysone occurs at similar levels in both sexes. Moreover, expression profiles of ecdysone response genes, ecdysone receptor and ecdysone-induced protein 75B, were also analyzed. Based on these expression profiles, we propose that the changes in ecdysteroid titer differ between males and females, and that high ecdysteroid titer is essential for directing male adult development., Competing Interests: The authors have declared that no competing interests exist.

Specimens of Stylops advarians were sampled by collecting foraging bees of Andrena milwaukeensis along the South Saskatchewan River within Saskatoon, Saskatchewan. As the foraging season progressed from early May till late June over three consecutive years (2016-2018), most stylopized bees possessed endoparasitic adult (neotenic) females of S. advarians protruding from the bee gaster's dorsum. In contrast, very few adult bees stylopized by male puparia, and no free-living males, were encountered. Over the sampling period, prevalence remained around 22% each year; mean intensity was 1.2 (range of 1-3 female parasites per bee); and parasite abundance was 0.27. Also newly reported for Stylops is the occurrence of one bee bearing four Stylops (two neotenic females and two males with puparia), plus another bee with a male puparium extruded from its gaster's sternites. Around 2 May each year, a high proportion of the earliest captured female bees were stylopized. However, non-stylopized female bees typically were not encountered until about 10 days later, suggesting the parasites manipulate female bee hosts to emerge earlier, in close synchrony to male bee emergence. First-instar larvae of S. advarians appeared from 22-25 May, indicating that adults of S. advarians matured and mated at similar times each season.

Systemic grass-endophytes of the genus Epichloë symbiotically infect the above-ground plant parts of many grass species, where they produce alkaloids in a grass- and endophyte-specific manner that are toxic or deterrent to herbivores. An increasing number of studies show cascading negative effects of endophyte-derived alkaloids that extend to higher trophic levels, harming beneficial insects, including those that control aphid populations. Lacewings are one of the major biological aphid controls, and are especially resistant to insecticides and pollutants, but their susceptibility to endophyte infection in the food chain has never been studied. Our study found variability in aphid population growth depending on the endophyte-grass chemotype, where aphid population growth was lowest on chemotypes known for producing high amounts of loline alkaloids. We also showed that larval and pupal development and mortality of the Common Green Lacewing (Chrysoperla carnea) was, in a non-choice experiment, not affected by endophyte infection in the food chain. This is a first indication that lacewings might be resistant to endophyte-derived alkaloids and could be robust biocontrol agents when applied together with endophyte-infected grass, possibly replacing chemical pesticides.

A remarkable new genus and species of Nemourinae, Sinonemura balangshana gen. et sp. n., is described from Balang Mountains, Sichuan, southwestern China. The description is based on morphology and molecular data. The Nemourinae genera related to the new taxon are re-evaluated on the basis of comparative functional morphology of male epiproct. Notes on the Asian distribution of the Nemourinae are also given., Competing Interests: The authors have declared that no competing interests exist.

Catalase is a widespread heme-containing enzyme, which converts hydrogen peroxide (H 2 O 2 ) to water and molecular oxygen, thereby protecting cells from the toxic effects of H 2 O 2 . Trypanosoma brucei is an aerobic protist, which conspicuously lacks this potent enzyme, present in virtually all organisms exposed to oxidative stress. To uncover the reasons for its absence in T. brucei, we overexpressed different catalases in procyclic and bloodstream stages of the parasite. The heterologous enzymes originated from the related insect-confined trypanosomatid Crithidia fasciculata and the human. While the trypanosomatid enzyme (cCAT) operates at low temperatures, its human homolog (hCAT) is adapted to the warm-blooded environment. Despite the presence of peroxisomal targeting signal in hCAT, both human and C. fasciculata catalases localized to the cytosol of T. brucei. Even though cCAT was efficiently expressed in both life cycle stages, the enzyme was active in the procyclic stage, increasing cell's resistance to the H 2 O 2 stress, yet its activity was suppressed in the cultured bloodstream stage. Surprisingly, following the expression of hCAT, the ability to establish the T. brucei infection in the tsetse fly midgut was compromised. In the mouse model, hCAT attenuated parasitemia and, consequently, increased the host's survival. Hence, we suggest that the activity of catalase in T. brucei is beneficial in vitro, yet it becomes detrimental for parasite's proliferation in both invertebrate and vertebrate hosts, leading to an inability to carry this, otherwise omnipresent, enzyme., (© 2019 Federation of European Biochemical Societies.)