Plant Chemical Defenses against Insect Herbivores—Using theWild Tobacco as a Model

The Nicotiana genus, commonly known as tobacco, holds significant importance as a crucial economic crop. Confrontedwith an abundance of herbivorous insects that pose a substantial threat to yield, tobacco has developed adiverse and sophisticated array of mechanisms, establishing itself as a model of plant ecological defense. Thisreview provides a concise overview of the current understanding of tobacco’s defense strategies against herbivores.Direct defenses, exemplified by its well-known tactic of secreting the alkaloid nicotine, serve as a potent toxinagainst a broad spectrum of herbivorous pests. Moreover, in response to herbivore attacks, tobacco enhancesthe discharge of volatile compounds, harnessing an indirect strategy that attracts the predators of the herbivores.The delicate balance between defense and growth leads to the initiation of most defense strategies only after aherbivore attack. Among plant hormones, notably jasmonic acid (JA), play central roles in coordinating thesedefense processes. JA signaling interacts with other plant hormone signaling pathways to facilitate the extensivetranscriptional and metabolic adjustments in plants following herbivore assault. By shedding light on these ecologicaldefense strategies, this review emphasizes not only tobacco’s remarkable adaptability in its natural habitatbut also offers insights beneficial for enhancing the resilience of current crops.

Global Change Effects on Plant Chemical Defenses against Insect Herbivores

This review focuses on individual effects of major global change factors, such as elevated CO2, O3, UV light and temperature, on plant secondary chemistry. These secondary metabolites are well-known for their role in plant defense against insect herbivory. Global change effects on secondary chemicals appear to be plant species-specifc and dependent on the chemical type. Even though plant chemical responses induced by these factors are highly variable, there seems to be some specificity in the response to different environmental stressors. For example, even though the production of phenolic compounds is enhanced by both elevated CO2 and UV light levels, the latter appears to primarily increase the concentrations of flavonoids. Likewise, specific phenolic metabolites seem to be induced by O3 but not by other factors, and an increase in volatile organic compounds has been particularly detected under elevated temperature. More information is needed regarding how global change factors influence inducibility of plant chemical defenses as well as how their indirect and direct effects impact insect performance and behavior, herbivory rates and pathogen attack. This knowledge is crucial to better understand how plants and their associated natural enemies will be affected in future changing environments.

Involvement of CYP347W1 in neurotoxin 3-nitropropionic acid-based chemical defense in mustard leaf beetle Phaedon cochleariae

Chrysomelina beetlesstore 3-nitropropionic acid in form of a pretoxin,isoxazolin-5-one glucoside-conjugated ester,to protect themselves against predators.Here we identified a cytochrome P450 monooxygenase,CYP347W1,to be involved in the production of the 3-nitropropionic acid moiety of the isoxazolin-5-one glucoside ester.Knocking down CYP347W1 led to a significant depletion in the concentration of the isoxazolin-5-one glucoside ester and an increase in the concentration of the isoxazolin-5-one glucoside in the larval hemolymph.Enzyme assays with the heterologously expressed CYP347W1 showed freeβ-alanine was not the direct substrate.Homology modeling indicated thatβ-alanine-CoA ester can fit into CYP347W1’s active site.Furthermore,we proved that Phaedon cochleariae eggs are not able to de novo synthesize 3-NPA,although both isoxazolin-5-one glucoside and its 3-NPA-conjugated ester are present in the eggs.These results provide direct evidence for the involvement of CYP347W1 in the biosynthesis of a P.cochleariae chemical defense compound.

Plant-herbivore assemblages under natural conditions are driven by plant size,not chemical defenses

Aims Plant secondary metabolites have been traditionally recognized as key traits regulating plant-herbivore assemblages.However,the ecological relevance of secondary metabolites as resistance mecha-nisms in comparison to other plant attributes,including physical,morphological or ecological traits,has been recently questioned.We aim to evaluate the role of chemical defenses,plant size and the presence of insect competitors on driving the differences in her-bivory damage under natural conditions.Methods We performed a replicated field study on the herbivore commu-nity associated with four Senecio species(S.lividus,S.vulgaris,S.inaequidens and S.pterophorus)during a full-reproductive season in Montseny Natural Park(catalonia,NE Spain).Pyrrolizidine alka-loids(PAs),the most characteristic chemical defenses of Senecio due to their toxic effects on herbivores,were analyzed by gas chro-matography.Individual plant size was estimated by the number of flower heads produced over the entire reproductive season.We used linear mixed models to explore the relationships between total PA concentrations,plant size and herbivory levels.Important Findings PA concentrations were not related to the natural guild of herbivores within any plant species or insect type.Moreover,no significant interactions were found between insect species sharing the same host plants.In contrast,herbivore abundance was positively related to plant size in S.vulgaris,S.lividus and S.inaequidens.We found no evidence that PAs confer an increased plant resistance against herbivores in Senecio.Our study supports the hypothesis that plant chemical defenses have a secondary role in determining plant-her-bivore assemblages in comparison to other plant traits under the complexity of natural conditions.

Effects of Exogenous Jasmonic Acid on Concentrations of Direct-Defense Chemicals and Expression of Related Genes in Bt (Bacillus thuringiensis) Corn (Zea mays)

Bt corn is one of the top three large-scale commercialized transgenic crops around the world. It is increasingly clear that the complementary durable approaches for pest control, which combine the endogenous defense of the crop with the introduced foreign genes, are promising alternative strategies for pest resistance management and the next generation of insect-resistant transgenic crops. In the present study, we tested the inducible effects of exogenous jasmonic acid （JA） on direct-defense chemical content, Bt protein concentration, and related gene expression in the leaves of Bt corn cultivar 34B24 and non-Bt cultivar 34B23 by chemical analysis, ELISA, and RT-PCR. The results show that the expression of LOX, PR-2a, MPI, and PR-I genes in the treated leaf （the first leaf） was promoted by exogenous JA both in 34B24 and 34B23. As compared with the control, the concentration of DIMBOA in the treated leaf was significantly increased by 63 and 18% for 34B24 and 34B23, respectively. The total phenolic acid was also increased by 24 and 12% for both 34B24 and 34B23. The Bt protein content of 34B24 in the treated leaf was increased by 13% but decreased significantly by 27% in the second leaf. The induced response of 34B24 was in a systemic way and was much stronger than that of 34B23. Those findings indicated that there is a synergistic interaction between Bt gene and internally induced chemical defense system triggered by externally applied JA in Bt corn.

Toxicity and Efficacy of Two Emulsifiable Concentrates of 2-Tridecanone against Red Imported Fire Ants

2-tridecanone is a major constitute of defensive secretion in tawny crazy ants, Nylanderia fulva that is an ant species reported to be able to displace red imported fire ants, Solenopsis invicta in the field. 2-tridecanone was proven to be toxic to S. invicta. Although 2-tridecanone is commercially available, utilization of this naturally occurring compound in fire ant management has received little attention. In this study, toxicity and efficacy of two 2-tridecanone formulations were assessed against S. invicta. Two emulsifiable concentrates were prepared using 2-tridecanone as an active ingredient, vegetable oil as a solvent and polyoxyethylene tridecyl ether as a surfactant. Piperonyl butoxide (PBO) was used as a synergist in one formulation. Both formulations showed significant toxicity in laboratory bioassays. In a field trial, at application rate of 5.28 mL/L and 14 days after mound drench treatment, 100% control was achieved for formulation with PBO and 90% control for the formulation without PBO. In conclusion, both formulations had significant efficacy against S. invicta. Due to the low mammalian toxicity of 2-tridecanone, no involvement of hazardous synthetic organic solvents, no phytotoxicity at applied concentrations, and relatively low cost, both formulations are promising alternatives to commercial insecticide products for fire ant mound drench. The outstanding efficacy of both formulations observed in this study warrants further research on their efficacy against other pest insects.

Inducible direct plant defense against insect herbivores： A review

Plants respond to insect herbivory with responses broadly known as direct defenses, indirect defenses, and tolerance. Direct defenses include all plant traits that affect susceptibility of host plants by themselves. Overall categories of direct plant defenses against insect herbivores include limiting food supply, reducing nutrient value, reducing preference, disrupting physical structures, and inhibiting chemical pathways of the attacking insect. Major known defense chemicals include plant secondary metabolites, protein inhibitors of insect digestive enzymes, proteases, lectins, amino acid deaminases and oxidases. Multiple factors with additive or even synergistic impact are usually involved in defense against a specific insect species, and factors of major importance to one insect species may only be of secondary importance or not effective at all against another insect species. Extensive qualitative and quantitative high throughput analyses of temporal and spatial variations in gene expression, protein level and activity, and metabolite concentration will accelerate not only the understanding of the overall mechanisms of direct defense, but also accelerate the identification of specific targets for enhancement of plant resistance for agriculture.

Increased reproductive capacity and physical defense but decreased tannin content in an invasive plant

Plant invasions create novel plant-insect interactions. The EICA （evolution of increased competitive ability） hypothesis proposes that invasive plants will reallocate resources from defense to growth and/or reproduction because they have escaped from their co-evolved insect natural enemies. Testing multiple herbivory by monophagous and oligophagous herbivores and simultaneous measurement of various plant traits will provide new insights into the evolutionary change ofinvasive plants. In this context, we conducted a common garden experiment to compare plant growth and reproduction, chemical and physical defense, and plant responses to herbivory by different types of herbivores between invasive North American populations and native East Asian populations of mile-a-minute weed, Persicariaperfoliata. We found that invasive mile-a-minnte exhibited lower biomass, flowered earlier and had greater reproductive output than plants from the native range. Compared with native populations, plants from invasive populations had lower tannin content, but exhibited higher prickle density on nodes and leaves. Thus our results partially support the EICA hypothesis. When exposed to the monophagous insect, Rhinoncomimus latipes and the oligophagous insects, Gallerucida grisescens and Smaragdina nigrifrons, more damage by herbivory was found on invasive plants than on natives. R. latipes, G. grisescens and S. nigrifrons had strong, moderate and weak impacts on the growth and reproduction of mile-a-minute, respectively. The results indicate that mile-a-minute may have evolved a higher reproductive capacity in the introduced range, and this along with a lack of oligophagous and monophagous herbivores in the new range may have contributed to its invasiveness in North America.

Defensive Sesquiterpenoids from Leaves of Eupatorium adenophorum

The invasive plant Eupatorium adenophorum Spreng has caused great economic loss in China, and is gravely threatening the native biodiversity and ecosystem. The plant has been phytochemically investigated for the defen- sive chemical substances in its leaves. Three active sesquiterpenoids were isolated and identified, which include a new sesquiterpenoid （1）, and two known sesquiterpenoids （2, 3）. Their structures were established by spectroscopic studies such as 1D- and 2D-NMR and MS analyses. Meanwhile, the antifeedant activities of these compounds against two generalist plant-feeding insects, Helicoverpa armigera and Spodoptera exigua, were carried out. Com- pound 1 showed significant antifeedant activity against S. exigua with ECs0 = 7.46 μg/cm^2, while compounds 2 and 3 were more active against H. armigera （EC50=2.57 and 3.04 μg/cm^2 respectively）. These findings suggest a de- fensive role of sesquiterpenoids in E. adenophorum against herbivores.