Changes in Bacterial Community Structure Across the Different Life Stages of Black Soldier Fly (Hermetia illucens).

The digestive capacity of organic compounds by the black soldier fly (BSF, Hermetia illucens, Diptera: Stratiomyidae, Linnaeus, 1758) is known to rely on complex larva-microbiota interactions. Although insect development is known to be a driver of changes of bacterial communities, the fluctuations along BSF life cycle in terms of composition and diversity of bacterial communities are still unknown. In this work, we used a metabarcoding approach to explore the differences in bacterial diversity along all four BSF developmental stages: eggs, larvae, pupae, and adult. We detected not only significant differences in bacterial community composition and species richness along the development of BSF, but also nine prevalent amplicon single variants (ASVs) forming the core microbiota. Out of the 2010 ASVs identified, 160 were significantly more abundant in one of the life stages. Moreover, using PICRUSt2, we inferred 27 potential metabolic pathways differentially used among the BSF life cycle. This distribution of metabolic pathways was congruent with the bacterial taxonomic distribution among life stages, demonstrating that the functional requirements of each phase of development are drivers of bacterial composition and diversity. This study provides a better understanding of the different metabolic processes occurring during BSF development and their links to changes in bacterial taxa. This information has important implications for improving bio-waste processing in such an economically important insect species.

Plant metabolism and defence strategies in the flowering stage: Time-dependent responses of leaves and flowers under attack.

Plants developing into the flowering stage undergo major physiological changes. Because flowers are reproductive tissues and resource sinks, strategies to defend them may differ from those for leaves. Thus, this study investigates the defences of flowering plants by assessing processes that sustain resistance (constitutive and induced) and tolerance to attack. We exposed the annual plant Brassica nigra to three distinct floral attackers (caterpillar, aphid and bacterial pathogen) and measured whole-plant responses at 4, 8 and 12 days after the attack. We simultaneously analysed profiles of primary and secondary metabolites in leaves and inflorescences and measured dry biomass of roots, leaves and inflorescences as proxies of resource allocation and regrowth. Regardless of treatments, inflorescences contained 1.2 to 4 times higher concentrations of primary metabolites than leaves, and up to 7 times higher concentrations of glucosinolates, which highlights the plant's high investment of resources into inflorescences. No induction of glucosinolates was detected in inflorescences, but the attack transiently affected the total concentration of soluble sugars in both leaves and inflorescences. We conclude that B. nigra evolved high constitutive rather than inducible resistance to protect their flowers; plants additionally compensated for damage by attackers via the regrowth of reproductive parts. This strategy may be typical of annual plants.

Bacterial Long-Range Warfare: Aerial Killing of Legionella pneumophila by Pseudomonas fluorescens.

Legionella pneumophila, the causative agent of Legionnaires' disease, is mostly found in man-made water systems and is one of the most closely monitored waterborne pathogens. With the aim of finding natural ways to control waterborne pathogens and thus further reduce the impact of disinfection by-products on human health, some studies have demonstrated the ability of bacteria to kill Legionella through the production of secondary metabolites or antimicrobial compounds. Here, we describe an unexpected growth inhibition of L. pneumophila when exposed to a physically separated strain of Pseudomonas fluorescens, designated as MFE01. Most of the members of the Legionellaceae family are sensitive to the volatile substances emitted by MFE01, unlike other bacteria tested. Using headspace solid-phase microextraction GC-MS strategy, a volatilome comparison revealed that emission of 1-undecene, 2-undecanone, and 2-tridecanone were mainly reduced in a Tn5-transposon mutant unable to inhibit at distance the growth of L. pneumophila strain Lens. We showed that 1-undecene was mainly responsible for the inhibition at distance in vitro, and led to cell lysis in small amounts, as determined by gas chromatography-mass spectrometry (GC-MS). Collectively, our results provide new insights into the mode of action of bacterial volatiles and highlight them as potent anti-Legionella agents to focus research on novel strategies to fight legionellosis. IMPORTANCE Microbial volatile compounds are molecules whose activities are increasingly attracting the attention of researchers. Indeed, they can act as key compounds in long-distance intrakingdom and interkingdom communication, but also as antimicrobials in competition and predation. In fact, most studies to date have focused on their antifungal activities and only a few have reported on their antibacterial properties. Here, we describe that 1-undecene, naturally produced by P. fluorescens, is a volatile with potent activity against bacteria of the genus Legionella. In small amounts, it is capable of inducing cell lysis even when the producing strain is physically separated from the target. This is the first time that such activity is described. This molecule could therefore constitute an efficient compound to counter bacterial pathogens whose treatment may fail, particularly in pulmonary diseases. Indeed, inhalation of these volatiles should be considered as a possible route of therapy in addition to antibiotic treatment.

Multiple Attack to Inflorescences of an Annual Plant Does Not Interfere with the Attraction of Parasitoids and Pollinators.

Plants in the flowering stage need to ensure reproduction by protecting themselves from attack and by preserving interactions with mutualist pollinators. When different plant mutualists are using the same type of cues, such as volatile compounds, attraction of parasitoids and pollinators may trade off. To explore this, we compared volatile emission of Brassica nigra plants in response to single or dual attack on their inflorescences. Additionally, we recorded flower visitation by pollinators and the attraction of parasitoids in the greenhouse and/or field. Brassica nigra were exposed in the flowering stage to one or two of the following three attackers: Brevicoryne brassicae aphids, Pieris brassicae caterpillars, and Xanthomonas campestris pv. raphani bacteria. We found that single attack by caterpillars, and dual attack by caterpillars plus aphids, induced the strongest changes in plant volatile emission. The caterpillars' parasitoid C. glomerata did not exhibit preference for plants exposed to caterpillars only vs. plants exposed to caterpillars plus aphids or plus bacteria. However, the composition of the pollinator community associated with flowers of B. nigra was affected by plant exposure to the attackers, but the total number of pollinators visiting the plants did not change upon attack. We conclude that, when B. nigra were exposed to single or dual attack on their inflorescences, the plants maintained interactions with natural enemies of the insect attackers and with pollinators. We discuss how chemical diversity may contribute to plant resilience upon attack.

Encuesta entomológica y caracterización de las viviendas con el vector de la enfermedad de Chagas, en aldea Chuchuapa, Santa Rosa, Guatemala / Entomological survey and characterization of houses with the vector of Chagas disease, in Chuchuapa village, Santa Rosa, Guatemala

En Guatemala en el 2015, el Ministerio de Salud Pública y Asistencia Social (MSPAS) reportó nuevos casos de la enfermedad de Chagas en la región suroriental del país, siendo Santa Rosa uno de los departamentos endémicos en donde Triatoma dimidiata es el principal transmisor. Se estimó la infestación de T. dimidiata en la aldea Chuchuapa, municipio de Santa María Ixhuatán, de abril a junio de 2019, se calcularon los índices en-tomológicos y se caracterizaron intra y peridomiciliar 149 viviendas (79.67% de las viviendas habitadas). Los datos se obtuvieron mediante un cuestionario mixto, en entrevista cara a cara y evaluación de las viviendas, por un equipo profesional experimentado, mediante el método y protocolo estándar hombre-hora. Se capturaron 20 triatominos en 10 viviendas, con un índice de infestación de 6.71% (10/149), un índice de densidad de 13.42% (20/149), 19 triatominos intradomiciliares y uno peridomiciliar, con un índice de infección natural por Tr y pano-soma cruzi intradomiciliar de 26.3% (5/19), índice de infección natural por T. cr u z i peridomiciliar de 100.0% (1/1) e índice de infección natural por T. c r u z i en los triatominos capturados de 30.0 % (6/20). Las viviendas infestadas presentaron techo de lámina, pared de bajareque con grietas y sin repello, piso de tierra y afiches en la pared, acumulación de leña u otros materiales tanto intra como peridomiciliar, convivencia con animales domésticos y tenencia de animales de corral. Los resultados representan un alto riesgo para la transmisión de la enfermedad de Chagas entre los habitantes de la aldea.

In Guatemala in 2015, the Ministry of Public Health and Social Assistance reported new cases of Chagas disease in the southeastern region of the country, with Santa Rosa being one of the endemic departments where Tr ia-toma dimidiata is the main transmitter. The infestation of T. dimidiata was estimated in the Chuchuapa village, municipality of Santa María Ixhuatán, from April to June 2019, the entomological indices were calculated and 149 dwellings were characterized intradomiciliary and peridomiciliary (79.67% of the inhabited dwellings). The data were obtained through a mixed questionnaire, in a face-to-face interview and evaluation of the dwellings, by an experienced professional team, using the standard man-hour method and protocol. 20 triatomines were captured in 10 homes, with an infestation index of 6.71% (10/149), a density index of 13.42% (20/149), 19 intradomiciliary triatomines and one peridomiciliary, with a natural infection index by intradomiciliary Trypanosomacruzi of 26.3% (5/19), natural infection index by peridomiciliary T. c r u z i of 100.0% (1/1) and natural infection index by T. c r u z i in captured triatomines of 30.0% (6/20). The infested dwellings have a sheet roof, awattle and daub wall with cracks and no wall plaster, dirt and posters on the wall, accumulation of firewood and other materials in both intradomiciliary and peridomiciliary, living with domestic animals and keeping poultry animals. The results represent a high risk for the transmission of Chagas disease among the villager.

The Evolution of Endophagy in Herbivorous Insects.

Herbivorous feeding inside plant tissues, or endophagy, is a common lifestyle across Insecta, and occurs in insect taxa that bore, roll, tie, mine, gall, or otherwise modify plant tissues so that the tissues surround the insects while they are feeding. Some researchers have developed hypotheses to explain the adaptive significance of certain endophytic lifestyles (e.g., miners or gallers), but we are unaware of previous efforts to broadly characterize the adaptive significance of endophagy more generally. To fill this knowledge gap, we characterized the limited set of evolutionary selection pressures that could have encouraged phytophagous insects to feed inside plants, and then consider how these factors align with evidence for endophagy in the evolutionary history of orders of herbivorous insects. Reviewing the occurrence of endophytic taxa of various feeding guilds reveals that the pattern of evolution of endophagy varies strongly among insect orders, in some cases being an ancestral trait (e.g., Coleoptera and Lepidoptera) while being more derived in others (e.g., Diptera). Despite the large diversity of endophagous lifestyles and evolutionary trajectories that have led to endophagy in insects, our consideration of selection pressures leads us to hypothesize that nutritionally based factors may have had a stronger influence on evolution of endophagy than other factors, but that competition, water conservation, and natural enemies may have played significant roles in the development of endophagy.

Gall-Inducing Parasites: Convergent and Conserved Strategies of Plant Manipulation by Insects and Nematodes.

Gall-inducing insects and nematodes engage in sophisticated interactions with their host plants. These parasites can induce major morphological and physiological changes in host roots, leaves, and other tissues. Sedentary endoparasitic nematodes, root-knot and cyst nematodes in particular, as well as gall-inducing and leaf-mining insects, manipulate plant development to form unique organs that provide them with food from feeding cells. Sometimes, infected tissues may undergo a developmental switch resulting in the formation of aberrant and spectacular structures (clubs or galls). We describe here the complex interactions between these plant-reprogramming sedentary endoparasites and their infected hosts, focusing on similarities between strategies of plant manipulation. We highlight progress in our understanding of the host plant response to infection and focus on the nematode and insect molecules secreted in planta. We suggest thatlooking at similarities may identify convergent and conserved strategies and shed light on the promise they hold for the development of new management strategies in agriculture and forestry.

Symbiosis disruption in the olive fruit fly, Bactrocera oleae (Rossi), as a potential tool for sustainable control.

BACKGROUND: The olive fruit fly Bactrocera oleae (Rossi) (OLF) is a major agricultural pest, whose control primarily relies on the use of chemical insecticides. Therefore, development of sustainable control strategies is highly desirable. The primary endosymbiotic bacterium of OLF, 'Candidatus Erwinia dacicola', is essential for successful larval development in unripe olive fruits. Therefore, targeting this endosymbiont with antimicrobial compounds may result in OLF fitness reduction and may exert control on natural populations of OLF. RESULTS: Here, we evaluate the impact of compounds with antimicrobial activity on the OLF endosymbiont. Copper oxychloride (CO) and the fungal metabolite viridiol (Vi), produced by Trichoderma spp., were used. Laboratory bioassays were carried out to assess the effect of oral administration of these compounds on OLF fitness and molecular analyses (quantitative polymerase chain reaction) were conducted to measure the load of OLF-associated microorganisms in treated flies. CO and Vi were both able to disrupt the symbiotic association between OLF and its symbiotic bacteria, determining a significant reduction in the endosymbiont and gut microbiota load as well as a decrease in OLF fitness. CO had a direct negative effect on OLF adults. Conversely, exposure to Vi significantly undermined larval development of the treated female's progeny but did not show any toxicity in OLF adults. CONCLUSIONS: These results provide new insights into the symbiotic control of OLF and pave the way for the development of more sustainable strategies of pest control based on the use of natural compounds with antimicrobial activity. © 2020 Society of Chemical Industry.

Salivary proteins of Phloeomyzus passerinii, a plant-manipulating aphid, and their impact on early gene responses of susceptible and resistant poplar genotypes.

Successful plant colonization by parasites requires the circumvention of host defenses, and sometimes a reprogramming of host metabolism, mediated by effector molecules delivered into the host. Using transcriptomic and enzymatic approaches, we characterized salivary glands and saliva of Phloeomyzus passerinii, an aphid exhibiting an atypical feeding strategy. Plant responses to salivary extracts of P. passerinii and Myzus persicae were assessed with poplar protoplasts of a susceptible and a resistant genotype, and in a heterologous Arabidopsis system. We predict that P. passerinii secretes a highly peculiar saliva containing effectors potentially interfering with host defenses, biotic stress signaling and plant metabolism, notably phosphatidylinositol phosphate kinases which seemed specific to P. passerinii. Gene expression profiles indicated that salivary extracts of M. persicae markedly affected host defenses and biotic stress signaling, while salivary extracts of P. passerinii induced only weak responses. The effector-triggered susceptibility was characterized by downregulations of genes involved in cytokinin signaling and auxin homeostasis. This suggests that P. passerinii induces an intracellular accumulation of auxin in susceptible host genotypes, which is supported by histochemical assays in Arabidopsis. This might in turn affect biotic stress signaling and contribute to host tissue manipulation by the aphid.

Gall Wasp Transcriptomes Unravel Potential Effectors Involved in Molecular Dialogues With Oak and Rose.

To gain insight into wasp factors that might be involved in the initial induction of galls on woody plants, we performed high throughput (454) transcriptome analysis of ovaries and venom glands of two cynipid gall wasps, Biorhiza pallida and Diplolepis rosae, inducing galls on oak and rose, respectively. De novo assembled and annotated contigs were compared to sequences from phylogenetically related parasitoid wasps. The relative expression levels of contigs were estimated to identify the most expressed gene sequences in each tissue. We identify for the first time a set of maternally expressed gall wasp proteins potentially involved in the interaction with the plant. Some genes highly expressed in venom glands and ovaries may act to suppress early plant defense signaling. We also identify gall wasp cellulases that could be involved in observed local lysis of plant tissue following oviposition, and which may have been acquired from bacteria by horizontal gene transfer. We find no evidence of virus-related gene expression, in contrast to many non-cynipid parasitoid wasps. By exploring gall wasp effectors, this study is a first step toward understanding the molecular mechanisms underlying cynipid gall induction in woody plants, and the recent sequencing of oak and rose genomes will enable study of plant responses to these factors.

Origin of gall-inducing from leaf-mining in Caloptilia micromoths (Lepidoptera, Gracillariidae).

In insects, the gall-inducing life-style has evolved independently many times. Several evolutionary pathways leading to this lifestyle have been proposed. While there is compelling evidence supporting surface-feeders and stem-borers as ancestral states of insect gall-inducers, an evolutionary pathway from leaf-miners remains hypothetical. Here we explored this question by comparing the developmental processes of two micromoths, a gall-inducer Caloptilia cecidophora (Lep., Gracillariidae), and its non-gall-inducing relative C. ryukyuensis. Like other Caloptilia, the first and second instars of C. cecidophora are leaf-miners and the gall is initiated inside the leaf mine by the third instar, thus suggesting leaf-mining as an ancestral, plesiomorphic state in this case. This is the first example of an insect species switching from leaf-mining to gall-inducing during larval development. The first two leaf-mining instars of C. cecidophora exhibit an absence of growth and a reduced time duration compared to C. ryukyuensis. The shortening of the duration of leaf-mining stages is apparently compensated in C. cecidophora by a larger egg size than C. ryukyuensis, and an additional larval instar during the gall phase.

Inside the horn of plenty: Leaf-mining micromoth manipulates its host plant to obtain unending food provisioning.

Leaves represent the main resource for herbivorous insects and their performances are mainly a function of leaf nutritional quality. Two feeding strategies are known to optimize the exploitation of leaf resources: leaf-miners that selectively feed on tissues of high nutritional quality and gall-inducers that induce the development of a new tissue showing an enhanced nutritional value. Some leaf-miners are known to also manipulate their nutritional environment, but do not affect plant development. Cases of callus proliferation in leaf-mines have been reported, however, the direct role of the insect in the formation of additional plant cells and the nutritional function of this tissue have never been established. Using an experimental approach, we show that leaf-mining larvae of micromoth, Borboryctis euryae (Lepidoptera: Gracillariidae), that grow on Eurya japonica (Pentaphylacaceae), actively induce callus proliferation within their leaf-mine at the fourth instar. We experimentally demonstrated that, at this developmental stage, the larva feeds exclusively on this newly formed tissue and feeding of the tissue is essential for completing larval stage. Phenological census revealed considerable expansion and variation of fourth instar duration caused by the continuous production of callus. We propose here the "cornucopia" hypothesis which states that the newly produced callus induced by the leaf-mining larvae provides virtually unending nourishment, which in turn allows flexible larval development time. This represents the first example of a leaf-miner manipulating plant development to its benefit, like a gall-inducer. We propose to name this life style "mine-galler".

A host-feeding wasp shares several features of nitrogen management with blood-feeding mosquitoes.

Adult feeding on hosts is common among parasitic wasps. The ingested host fluid is rich in nutrients, especially proteins. A study on Eupelmus vuilleti (Hymenoptera: Eupelmidae), a host-feeding parasitoid of larvae of Callosobruchus maculatus (F.) (Coleoptera: Bruchidae), showed that the carbohydrates (maybe lipids) but not proteins, gained from host feeding accounted for the increased egg production. Thus, host protein is probably utilized for general adult metabolism, allowing conservation of carbohydrate and/or lipid resources for direct allocation to oocytes. In that case, there should be increased N excretion by female parasitoids. To test this, we studied the dynamics of excretion in E. vuilleti with and without host exposure. The aim of this work was threefold: (i) to identify the major N-containing compounds in adult excreta, (ii) to assess whether protein consumption during host feeding increased the amount of N excreted, and (iii), if so, to compare the increase in N excreted with the amount taken in during a single host feeding. We found that uric acid is the predominant N-containing metabolite in excreta, although small quantities of urea and traces of allantoin were also found. A calculation of the N budget showed that the extra quantity of N excreted following a host meal corresponds to the quantity ingested, confirming that host-feeding in this species offers little or no net quantitative benefit in N allocation to oocytes, although the allocation of specific amino acids from host feeding cannot be discounted. Interestingly, host-feeding in parasitoids appears analogous to vertebrate blood-feeding in mosquitoes, both in terms of the N-containing compounds excreted and the offset of acquired N to metabolism, rather than to oocytes. Further comparative and detailed investigations of N excretion in insects living on other N-rich fluids might establish further metabolic commonalities.

Caterpillars induce jasmonates in flowers and alter plant responses to a second attacker.

In nature, herbivorous insects and plant pathogens are generally abundant when plants are flowering. Thus, plants face a diversity of attackers during their reproductive phase. Plant responses to one attacker can interfere with responses to a second attacker, and phytohormones that orchestrate plant reproduction are also involved in resistance to insect and pathogen attack. We quantified phytohormonal responses of flowering plants exposed to single or dual attack and studied resistance mechanisms of plants in the flowering stage. Flowering Brassica nigra were exposed to either a chewing caterpillar, a phloem-feeding aphid or a bacterial pathogen, and plant hormonal responses were compared with dual attack situations. We quantified phytohormones in inflorescences and leaves, and determined the consequences of hormonal changes for components of direct and indirect plant resistance. Caterpillars were the main inducers of jasmonates in inflorescences, and the phytohormonal profile of leaves was not affected by either insect or pathogen attack. Dual attack increased plant resistance to caterpillars, but compromised resistance to aphids. Parasitoid performance was negatively correlated with the performance of their hosts. We conclude that plants prioritize resistance of reproductive tissues over vegetative tissues, and that a chewing herbivore species is the main driver of responses in flowering B. nigra.

CHASE-Containing Histidine Kinase Receptors in Apple Tree: From a Common Receptor Structure to Divergent Cytokinin Binding Properties and Specific Functions.

Cytokinin signaling is a key regulatory pathway of many aspects in plant development and environmental stresses. Herein, we initiated the identification and functional characterization of the five CHASE-containing histidine kinases (CHK) in the economically important Malus domestica species. These cytokinin receptors named MdCHK2, MdCHK3a/MdCHK3b, and MdCHK4a/MdCHK4b by homology with Arabidopsis AHK clearly displayed three distinct profiles. The three groups exhibited architectural variations, especially in the N-terminal part including the cytokinin sensing domain. Using a yeast complementation assay, we showed that MdCHK2 perceives a broad spectrum of cytokinins with a substantial sensitivity whereas both MdCHK4 homologs exhibit a narrow spectrum. Both MdCHK3 homologs perceived some cytokinins but surprisingly they exhibited a basal constitutive activity. Interaction studies revealed that MdCHK2, MdCHK4a, and MdCHK4b homodimerized whereas MdCHK3a and MdCHK3b did not. Finally, qPCR analysis and bioinformatics approach pointed out contrasted expression patterns among the three MdCHK groups as well as distinct sets of co-expressed genes. Our study characterized for the first time the five cytokinin receptors in apple tree and provided a framework for their further functional studies.

Maternal age affects offspring nutrient dynamics.

The internal physiological state of a mother can have major effects on her fitness and that of her offspring. We show that maternal effects in the parasitic wasp Eupelmus vuilleti become apparent when old mothers provision their eggs with less protein, sugar and lipid. Feeding from a host after hatching allows the offspring of old mothers to overcome initial shortages in sugars and lipids, but adult offspring of old mothers still emerged with lower protein and glycogen quantities. Reduced egg provisioning by old mothers had adverse consequences for the nutrient composition of adult female offspring, despite larval feeding from a high-quality host. Lower resource availability in adult offspring of old mothers can affect behavioural decisions, life histories and performance. Maternal effects on egg nutrient provisioning may thus affect nutrient availability and fitness of future generations in oviparous animals.

Dynamics and origin of cytokinins involved in plant manipulation by a leaf-mining insect.

Several herbivorous insects and plant-associated microorganisms control the phytohormonal balance, thus enabling them to successfully exploit the plant by inhibiting plant defenses and withdrawing plant resources for their own benefit. The leaf-mining moth Phyllonorycter blancardella modifies the cytokinin (CK) profile of mined leaf-tissues, and the insect symbiotic bacteria Wolbachia is involved in the plant manipulation to the benefit of the insect host. To gain a deeper understanding into the possible origin and dynamics of CKs, we conducted an extensive characterization of CKs in larvae and in infected apple leaves. Our results show the enhanced CK levels in mines, both on green and yellow leaves, allowing insects to control their nutritional supply under fluctuating environmental conditions. The spatial distribution of CKs within the mined leaves shows that hormone manipulation is strictly limited to the mine suggesting the absence of CK translocation from distant leaf areas toward the insect feeding site. Mass spectrometry analyses reveal that major CK types accumulating in mines and larvae are similar to what is observed for most gall-inducers, suggesting that strategies underlying the plant manipulation may be shared between herbivorous insects with distinct life histories. Results further show that CKs are detected in the highest levels in larvae, reinforcing our hypothesis that CKs accumulating in the mines originate from the insect itself. Presence of bacteria-specific methylthio-CKs is consistent with previous results suggesting that insect bacterial symbionts contribute to the observed phenotype. Our study provides key findings toward the understanding of molecular mechanisms underlying this intricate plant-insect-microbe interaction.

Insect-induced effects on plants and possible effectors used by galling and leaf-mining insects to manipulate their host-plant.

Gall-inducing insects are iconic examples in the manipulation and reprogramming of plant development, inducing spectacular morphological and physiological changes of host-plant tissues within which the insect feeds and grows. Despite decades of research, effectors involved in gall induction and basic mechanisms of gall formation remain unknown. Recent research suggests that some aspects of the plant manipulation shown by gall-inducers may be shared with other insect herbivorous life histories. Here, we illustrate similarities and contrasts by reviewing current knowledge of metabolic and morphological effects induced on plants by gall-inducing and leaf-mining insects, and ask whether leaf-miners can also be considered to be plant reprogrammers. We review key plant functions targeted by various plant reprogrammers, including plant-manipulating insects and nematodes, and functionally characterize insect herbivore-derived effectors to provide a broader understanding of possible mechanisms used in host-plant manipulation. Consequences of plant reprogramming in terms of ecology, coevolution and diversification of plant-manipulating insects are also discussed.