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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.)

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.)

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.)

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.)

Insect courtship parades consist of series of innate and stereotyped behaviours that become hardwired-in during the development of the nervous system. As such, insect courtship behaviour provides an excellent model for probing the principles of neuronal assembly, which underlie patterns of behaviour. Here, we present the main advances of recent studies - in species all the way from flies to planthoppers - and we envisage how these could lead to further propitious findings., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Insect herbivores express tremendous ontogenetic variation in traits related to growth and maturation, but also as an evolutionary consequence of ecological interactions with plants and predators. These selective pressures can either reinforce or restrict expression of particular ontogenetic strategies, allowing herbivores to simultaneously cope with plant resistance and risk of predation through ontogenetic change. For example, whereas an increase in defense-sabotaging behavior, aposematism and sequestration along herbivore ontogeny seems to be reinforced by both bottom-up and top-down forces, some ontogenetic trends in anti-predator behavior can be limited by plant resistance. Communication among plants, herbivores and their natural enemies is also influenced by insect ontogenies. The study of ontogenetic strategies of herbivores requires the assessment of the genetic variation, heritability and adaptive value across herbivore development, considering the variation in plant quality and predation risk., (Copyright © 2018 Elsevier Inc. All rights reserved.)

The physiological control of reproduction in insects depends on a combination of environmental and internal cues. In the adult stage, insects become sexually mature and generate gametes. In females, the latter process is designated as oogenesis. Peptides are a versatile class of extracellular signalling molecules that regulate many processes, including oogenesis. At present, the best documented physiological control mechanism of insect oogenesis is the insulin-related peptide signalling pathway. It regulates different stages of the process and provides a functional link between nutritional status and reproduction. Several other peptides have been shown to exert gonadoregulatory activities, but in most cases their exact mode of action still has to be unravelled and their effects on oogenesis could be direct or indirect. Some regulatory peptides, such as the Drosophila sex peptide, are being transferred from the male to the female during the mating process., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Juvenile hormone (JH) plays a crucial role in insect reproduction, but its molecular mode of action only became clear within the last decade. We here review recent findings revealing the intricate crosstalk between JH and ecdysone signaling with nutrient sensing pathways in Drosophila melanogaster, Aedes aegypti, Tribolium castaneum and Locusta migratoria. The finding for a critical role of ecdysis triggering hormone (ETH) in both molting and ooogenesis now also highlights the importance of an integrated view of development and reproduction. Furthermore, insights from non-model insects, especially so social Hymenoptera and termites, where JH function gradually becomes decoupled from reproduction and plays a role in division of labor, emphasize the need to consider life cycle and life history strategies when studying insect reproductive physiology., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Ontogeny of animal personality is still an open question. Testing whether personality traits correlated with state variables (e.g. metabolic rate, hormones) and/or life history traits, and which ones are involved, requires more empirical studies. Insects with metamorphosis represent a good opportunity to tackle this question. Because of the various degrees of internal (physiological, nervous) and environmental changes linked to metamorphosis they allow testing whether these modifications drive consistency in personality traits between immature and adult stages. In this review, we establish general predictions for the effects of metamorphosis on personality in insects with complete or incomplete metamorphosis and suffering from a strong or weak niche shift after metamorphosis. We then reviewed the still rare empirical literature and discuss future research axes., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Developmental plasticity is a key driver of the extraordinary ecological success of insects. Epigenetic mechanisms provide an important link between the external stimuli that initiate polyphenisms, and the stable changes in gene expression that govern alternative insect morphs. We review the epigenetics of orthopteroid insects, focussing on recent research on locusts and termites, two groups which display high levels of phenotypic plasticity, and for which genome sequences have become available in recent years. We examine research on the potential role of DNA methylation, histone modifications, and non-coding RNAs in the regulation of gene expression in these insects. DNA methylation patterns in orthopteroids share a number of characteristics with those of hymenopteran insects, although methylation levels are much higher, and extend to introns and repeat elements. Future examinations of epigenetic mechanisms in these insects will benefit from comparison of tissues from aged-matched individuals from alternative morphs, and adequate biological replication., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Many insects have the ability to develop alternative morphologies in response to specific environmental signals such as photoperiod, temperature, nutrition and crowding. These signals are integrated by the brain and result in alternative patterns of secretion of developmental hormones like ecdysone, juvenile hormone and insulin-like growth factors, which, in turn, direct alternative developmental trajectories. Insulin signaling appears to be particularly important when the polyphenism involves differences in the sizes of the body, appendages and other structures, such as wings, mandibles and horns. Here we review recent advances in understanding the role of insulin signaling, and its interaction with other hormones, in the development of polyphenisms., (Copyright © 2017. Published by Elsevier Inc.)

Differential growth, the phenomenon where parts of the body grow at different rates, is necessary to generate the complex morphologies of most multicellular organisms. Despite this central importance, how differential growth is regulated remains largely unknown. Recent discoveries, particularly in insects, have started to uncover the molecular-genetic and physiological mechanisms that coordinate growth among different tissues throughout the body and regulate relative growth. These discoveries suggest that growth is coordinated by a network of signals that emanate from growing tissues and central endocrine organs. Here we review these findings and discuss their implications for understanding the regulation of relative growth and the evolution of morphology., (Copyright © 2017. Published by Elsevier Inc.)

Insects show impressive diversity in adult body size across species, and within species adult body size is sensitive to numerous environmental conditions, particularly to changes in nutrition. Body size in adult insects correlates with a number of important fitness-related traits such as fecundity, longevity, stress resistance, and mating success. Over the past few decades, the field of insect body size regulation has made impressive progress towards understanding the signalling pathways that regulate body size in response to nutrition. These studies have shown that conserved nutrition-sensitive signalling pathways act in animals from insects to vertebrates to regulate growth. In particular, pathways like the insulin/insulin-like growth factor signalling (IIS) pathway and the Target of rapamycin (TOR) pathway respond to the levels of dietary nutrients to adjust both the rate of growth and the duration of the growth period. They do this not only by regulating organ growth, but also by modifying the rates of synthesis and circulating concentrations of key developmental hormones. Although the mechanisms through which this occurs have been well documented in one insect, the fruit fly Drosophila melanogaster, it is becoming increasingly clear that the downstream mechanisms through which IIS and TOR signalling alter size in response to nutrition differ between organs and across species. In this review, we highlight how understanding the organ-specific effects of IIS/TOR signalling are key to revealing the diversity of size control mechanisms across insects., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Insects show a tremendous morphological variety and have been a subject of studying morphological evolution. In legs, the tarsus is especially variable in the number of subsegments (tarsal segments) and their proportion unlike other leg segments. Recent studies in Drosophila melanogaster have revealed details of the tarsal development: regionalization of the tarsal region through integration of regulatory network and its growth, determination of the joint-forming region in each segment through strict regulation of Notch activity, changes in tissue morphology through regulation of RhoGTPases regulators and localized cell death, and finally, the morphogenetic mechanism of the ball-and-socket joint between tarsal segments. The substantial knowledge of the tarsal development makes it a suitable model for studying mechanisms of morphological evolution and diversification., (Copyright © 2016 Elsevier Inc. All rights reserved.)

The wide variety of external morphologies has underlain the evolutionary success of insects. The insect exoskeleton, or cuticle, which covers the entire body and constitutes the external morphology, is extracellular matrix produced by the epidermis. How is cuticle shaped during development? Past studies have mainly focused on patterning, differentiation and morphogenesis of the epidermis. Recently, however, it is becoming clear that cuticle itself plays important and active roles in regulation of cuticle shape. Studies in the past several years show that pre-existing cuticle can influence shaping of new cuticle, and cuticle can sculpt its own shape through its material property. In this review, I summarize recent advances and discuss future prospects., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Secondary sexual traits are those traits other than the primary gametes that distinguish the sexes of a species. The development of secondary sexual traits occurs when sexually dimorphic factors, that is, molecules differentially produced by primary sex determination systems in males and females, are integrated into the gene regulatory networks responsible for sexual trait development. In insects, these molecular asymmetric factors were always considered to originate inside the trait-building cells, but recent work points to external factors, such as hormones, as potential candidates mediating secondary sexual trait development. Here, we review examples of the different molecular mechanisms producing sexually dimorphic traits in insects, and suggest a need to revise our understanding of secondary sexual trait development within the insect lineage., (Copyright © 2016 Elsevier Inc. All rights reserved.)