Lepidopteran insects are one of the most widespread and speciose lineages on Earth, with many common pests and beneficial insect species. The evolutionary success of their diversification depends on the essential functions of gut microorganisms. This diverse gut microbiota of lepidopteran insects provides benefits in nutrition and reproductive regulation and plays an important role in the defence against pathogens, enhancing host immune homeostasis. In addition, gut symbionts have shown promising applications in the development of novel tools for biological control, biodegradation of waste, and blocking the transmission of insect-borne diseases. Even though most microbial symbionts are unculturable, the rapidly expanding catalogue of microbial genomes and the application of modern genetic techniques offer a viable alternative for studying these microbes. Here, we discuss the gut structure and microbial diversity of lepidopteran insects, as well as advances in the understanding of symbiotic relationships and interactions between hosts and symbionts. Furthermore, we provide an overview of the function of the gut microbiota, including in host nutrition and metabolism, immune defence, and potential mechanisms of detoxification. Due to the relevance of lepidopteran pests in agricultural production, it can be expected that the research on the interactions between lepidopteran insects and their gut microbiota will be used for biological pest control and protection of beneficial insects in the future.

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Diversity and Functional Roles of the Gut Microbiota in Lepidopteran Insects

With the increase in world population, the demography of humans is estimated to be exceeded and it has become a major challenge to provide an adequate amount of food, feed, and agricultural products majorly in developing countries. The use of chemical fertilizers causes the plant to grow efficiently and rapidly to meet the food demand. The drawbacks of using a higher quantity of chemical or synthetic fertilizers are environmental pollution, persistent changes in the soil ecology, physiochemical composition, decreasing agricultural productivity and cause several health hazards. Climatic factors are responsible for enhancing abiotic stress on crops, resulting in reduced agricultural productivity. There are various types of abiotic and biotic stress factors like soil salinity, drought, wind, improper temperature, heavy metals, waterlogging, and different weeds and phytopathogens like bacteria, viruses, fungi, and nematodes which attack plants, reducing crop productivity and quality. There is a shift toward the use of biofertilizers due to all these facts, which provide nutrition through natural processes like zinc, potassium and phosphorus solubilization, nitrogen fixation, production of hormones, siderophore, various hydrolytic enzymes and protect the plant from different plant pathogens and stress conditions. They provide the nutrition in adequate amount that is sufficient for healthy crop development to fulfill the demand of the increasing population worldwide, eco-friendly and economically convenient. This review will focus on biofertilizers and their mechanisms of action, role in crop productivity and in biotic/abiotic stress tolerance.

Overview of biofertilizers in crop production and stress management for sustainable agriculture

Although the use of synthetic chemicals is the principal method for insect pest management, their widespread application has led to numerous side effects, including environmental pollution and threats to human and animal health. Plant essential oils have been introduced as promising natural substitutes for synthetic insecticides. However, high volatility and/or low durability are the main limiting factors for essential oil application for control of insect pests. Accordingly, along with an evaluation of the fumigant toxicity of Eucalyptus largiflorens essential oil against the cowpea weevil, Callosobruchus maculatus, essential oil was nanoencapsulated by two mesoporous silicates, MCM-41 and zeolite 3A, to enhance fumigant persistence and toxicity. The chemical profile of essential oil was also analyzed through gas chromatographic-mass spectrometry. E. largiflorens essential oil showed significant concentration-dependent toxicity against insect pests; a concentration of 5.16 μL/L resulted in 100% mortality after 48 h. The toxicity of essential oil could be attributed to the presence of various insecticidal terpenes, such as spathulenol (15.6%), cryptone (7.0%), and 1,8-cineole (5.8%). Fumigant persistence was increased from 6 days to 19 and 17 days for pure and capsulated essential oil with MCM-41 and Zeolite 3A, respectively. The insect mortality also increased from 99 insects in pure essential oil to 178 and 180 insects in MCM-41 and Zeolite 3A encapsulated formulations, respectively. Therefore, the encapsulation of E. largiflorens essential oil by MCM- 41 and Zeolite 3A is a beneficial method for enhancing its persistence and toxicity against C. maculatus.

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Encapsulation of Eucalyptus largiflorens Essential Oil by Mesoporous Silicates for Effective Control of the Cowpea Weevil, Callosobruchus maculatus (Fabricius) (Coleoptera: Chrysomelidae)

Protease Inhibitors: An Induced Plant Defense Mechanism Against Herbivores

Pod borer Helicoverpa armigera, a polyphagus herbivorous pest, tremendously incurs crop damage in economically important crops. This necessitates the identification and utility of novel genes for the control of the herbivore. The present study deals with the characterization of a flavonoid 3′5′ hydroxylase_2 (F3′5′H_2) from a pigeonpea wild relative Cajanus platycarpus, possessing a robust chemical resistance response to H. armigera. Though F3′5′H_2 displayed a dynamic expression pattern in both C. platycarpus (Cp) and the cultivated pigeonpea, Cajanus cajan (Cc) during continued herbivory, CpF3′5′H_2 showed a 4.6-fold increase vis a vis 3-fold in CcF3′5′H_2. Despite similar gene copy numbers in the two Cajanus spp., interesting genic and promoter sequence changes highlighted the stress responsiveness of CpF3′5′H_2. The relevance of CpF3′5′H_2 in H. armigera resistance was further validated in CpF3′5′H_2-overexpressed transgenic tobacco based on reduced leaf damage and increased larval mortality through an in vitro bioassay. As exciting maiden clues, CpF3′5′H_2 deterred herbivory in transgenic tobacco by increasing total flavonoids, polyphenols and reactive oxygen species (ROS) scavenging capacity. To the best of our knowledge, this is a maiden attempt ascertaining the role of F3′5′H_2 gene in the management of H. armigera. These interesting leads suggest the potential of this pivotal branch-point gene in biotic stress management programs.

https://www.mdpi.com/1422-0067/24/2/1755/pdf?version=1673854452

Cajanus platycarpus Flavonoid 3′5′ Hydroxylase_2 (CpF3′5′H_2) Confers Resistance to Helicoverpa armigera by Modulating Total Polyphenols and Flavonoids in Transgenic Tobacco

Plants have evolved several adaptive strategies through physiological changes in response to herbivore attacks. Plant secondary metabolites (PSMs) are synthesized to provide defensive functions and regulate defense signaling pathways to safeguard plants against herbivores. Herbivore injury initiates complex reactions which ultimately lead to synthesis and accumulation of PSMs. The biosynthesis of these metabolites is regulated by the interplay of signaling molecules comprising phytohormones. Plant volatile metabolites are released upon herbivore attack and are capable of directly inducing or priming hormonal defense signaling pathways. Secondary metabolites enable plants to quickly detect herbivore attacks and respond in a timely way in a rapidly changing scenario of pest and environment. Several studies have suggested that the potential for adaptation and/or resistance by insect herbivores to secondary metabolites is limited. These metabolites cause direct toxicity to insect pests, stimulate antixenosis mechanisms in plants to insect herbivores, and, by recruiting herbivore natural enemies, indirectly protect the plants. Herbivores adapt to secondary metabolites by the up/down regulation of sensory genes, and sequestration or detoxification of toxic metabolites. PSMs modulate multi-trophic interactions involving host plants, herbivores, natural enemies and pollinators. Although the role of secondary metabolites in plant-pollinator interplay has been little explored, several reports suggest that both plants and pollinators are mutually benefited. Molecular insights into the regulatory proteins and genes involved in the biosynthesis of secondary metabolites will pave the way for the metabolic engineering of biosynthetic pathway intermediates for improving plant tolerance to herbivores. This review throws light on the role of PSMs in modulating multi-trophic interactions, contributing to the knowledge of plant-herbivore interactions to enable their management in an eco-friendly and sustainable manner.

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Plant Secondary Metabolites as Defense Tools against Herbivores for Sustainable Crop Protection

Nutritionally rich cucumber seeds remain in demand in the agricultural, health and cosmetic sectors as they are essential for a successful crop stand establishment and seed-based products. However, the production of cucumber seeds is impeded by source limitation and nutrient deficiency. The foliar application of micronutrients can supplement this deficiency and overcome the physiological setback. An experiment was undertaken to compare the impacts of the foliar application of Fe and Zn, as nanoparticles and fertilizers, on the yield and seed quality of cucumber under open and protected environments. A foliar spray of nano-ZnO (ZnNPs) and nano-Fe3O4 (FeNPs) at 100, 200 and 300 mg L−1, as well as ZnSO4 and FeSO4 as fertilizer (0.5%), was conducted at the vegetative stage and pre- and post-flowering stages. The NPs had a greater efficacy in an open field than in the protected (naturally ventilated poly house) environment. The application of both NPs increased seed yield (51.7–52.2%), total chlorophyll content (15.9–17.3%) and concentration of Zn and Fe in the fruit and the seed, by 2.0–58.5% and 5.0–30.5%, respectively. A significant increase in starch, soluble proteins, soluble sugars and oil content was observed in the seeds from the NP treated plants. NP treatment also enhanced the germination-related parameters, such as percent germination (16.8–17.0%), rate of germination (18.0–22.2%) and seedling vigor (59.8–72.6%). The biochemical characterization showed a significant improvement in the seed water uptake and the activity of hydrolytic enzymes (amylase and protease) in the germinating seed. The involvement of reactive oxygen species (superoxide anion and hydrogen peroxide) and antioxidant enzymes (Superoxide dismutase, Catalase and Peroxidase) in the germination process was indicated by an increase in their activities in the seeds from NP treated plants. Hence, the study proposes the potential benefit of the foliar application of 300 mg L−1 ZnNPs and 200 mg L−1 FeNPs at crucial stages of plant growth to improve the yield and seed quality in cucumbers.

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Impact of Foliar Application of ZnO and Fe3O4 Nanoparticles on Seed Yield and Physio-Biochemical Parameters of Cucumber (Cucumis sativus L.) Seed under Open Field and Protected Environment vis a vis during Seed Germination

Vikas K. Singh

Papers

Plant Secondary Metabolites as Defense Tools against Herbivores for Sustainable Crop Protection

Impact of Foliar Application of ZnO and Fe3O4 Nanoparticles on Seed Yield and Physio-Biochemical Parameters of Cucumber (Cucumis sativus L.) Seed under Open Field and Protected Environment vis a vis during Seed Germination