Terpenoids of plants in arid environments.

Arid and desert environments are characterized by the sparse and discontinuous vegetation cover. Species that have been able to survive difficult bioclimatic conditions and adapt from generation to generation in these areas had to develop physiological and biochemical mechanisms of tolerance and/or resistance. The use of secondary metabolites, specifically terpenoids, is predominant in most of the biotic and abiotic interactions in which these plants are involved. Studies have shown their roles in the prevention of oxidative stress by intervening in thermo-tolerance, water stress, and salt stress generalized in a model of "the protective role of volatile compounds" explained by a single biochemical mechanism. Other studies have proven the functions of terpenoids in direct and indirect defenses against natural enemies, herbivores, and pathogenic microorganisms, in the attraction of pollinators, in competition and facilitation and other interactions between plants. This review mainly summarizes the recent research progress on the adaptation mechanisms of plants in arid environments and the biological and ecological roles of terpenoids in the various biotic and abiotic interactions.

Effects of Prohydrojasmon on the Number of Infesting Herbivores and Biomass of Field-Grown Japanese Radish Plants

Prohydrojasmon (PDJ), an analog of jasmonic acid (JA), was found to induce direct and indirect defenses against herbivores in non-infested plants. To test whether PDJ can be used for pest control in crop production, we conducted experiments in pesticide-free Japanese radish fields from October 4 to December 12 in 2015. Twenty-four Japanese radish plants in three plots were treated with a 100 times-diluted commercial formulation (5%) of PDJ (treated plants), and 24 plants in three different plots were treated with water (control plants) until November 29 every week. Throughout the observation period, the number of aphids, leaf-mining fly larvae, vegetable weevils, and thrips was significantly lower on the treated plants than on the control plants. In contrast, the number of lepidopteran larvae was not significantly different between the treated and control plants throughout the study period. Parasitized aphids (mummies) were also observed in both plots. Poisson regression analyses showed that a significantly higher number of mummies was recorded on the treated plants as compared to that on the control plants when the number of aphids increased. This suggested that PDJ application to Japanese radish plants attracted more parasitoid wasps on the treated plants than on the control plants. We also identified eight terpenoids and methyl salicylate as the PDJ-induced plant volatiles in the headspace of the treated plants. Some of these volatiles might be responsible for attracting aphid-parasitoid wasps in the field. However, for other insect pests, we did not find any natural enemies. Interestingly, the genes of the JA and salicylic acid signaling pathways were differentially upregulated in the treated plants. We also observed that the PDJ treatments induced the expression of the genes related to glucosinolate biosynthesis and the subsequent isothiocyanate formation. Additionally, the weights of both the aboveground and belowground parts of the treated plants were significantly lower than those of the respective parts of the control plants. These results indicated that the treatment of Japanese radish plants with a 100 times-diluted commercial formulation of PDJ induced their direct and indirect defenses against several insect pest species to reduce their numbers, and negatively affected their biomass.

Bacterial Endophytes: The Hidden Actor in Plant Immune Responses against Biotic Stress

Bacterial endophytes constitute an essential part of the plant microbiome and are described to promote plant health by different mechanisms. The close interaction with the host leads to important changes in the physiology of the plant. Although beneficial bacteria use the same entrance strategies as bacterial pathogens to colonize and enter the inner plant tissues, the host develops strategies to select and allow the entrance to specific genera of bacteria. In addition, endophytes may modify their own genome to adapt or avoid the defense machinery of the host. The present review gives an overview about bacterial endophytes inhabiting the phytosphere, their diversity, and the interaction with the host. Direct and indirect defenses promoted by the plant–endophyte symbiont exert an important role in controlling plant defenses against different stresses, and here, more specifically, is discussed the role against biotic stress. Defenses that should be considered are the emission of volatiles or antibiotic compounds, but also the induction of basal defenses and boosting plant immunity by priming defenses. The primed defenses may encompass pathogenesis-related protein genes (PR family), antioxidant enzymes, or changes in the secondary metabolism.

The ecological consequences of herbivore-induced plant responses on plant–pollinator interactions

Plant induced responses to herbivory have long been found to function as plant direct and indirect defenses and to be major drivers of herbivore community and population dynamics. While induced defenses are generally understood as cost-saving strategies that allow plants to allocate valuable resources into defense expression, it recently became clear that, in particular, induced metabolic changes can come with significant ecological costs. In particular, interactions with mutualist pollinators can be significantly compromised by herbivore-induced changes in floral morphology and metabolism. We review recent findings on the evidence for ecological conflict between defending against herbivores and attracting pollinators while using similar modes of information transfer (e.g. visual, olfactory, tactile). Specifically, we discuss plant traits and mechanisms through which plants mediate interactions between antagonists and mutualist and present functional hypotheses for how plants can overcome the resulting conflicts.

Polyphenol-Rich Purple Corn Pericarp Extract Adversely Impacts Herbivore Growth and Development

Plant secondary metabolites such as terpenes, phenolics, glycosides, and alkaloids play various functional roles including pigmentation, foliar and floral volatile synthesis, hormonal regulation, and direct and indirect defenses. Among these, phenolic compounds are commonly found in plants, but vary in the distribution of their specific compounds among plant families. Polyphenols, including anthocyanins and tannins, are widely distributed and have been well documented for their roles- primarily in plant pigmentation and also in plant defenses. However, commercialization of such compounds for use in insect pest management is severely hampered by expensive, inefficient, and time-consuming extraction protocols. Using a recently developed inexpensive and easy extraction method using the byproducts of pigmented (purple) corn processing, we examined whether the crude pericarp extract rich in polyphenols can affect the growth and development of tobacco hornworm (Manduca sexta L.) caterpillars. Our findings show that purple corn pericarp extract negatively affected M. sexta egg hatching and larval mass gain and prolonged developmental time compared to regular yellow corn extract or an artificial control diet. We also found that these effects were more severe during the early stages of caterpillar development. These results conclusively demonstrate that purple corn pericarp, an inexpensive by-product of the corn milling industry, is a valuable product with excellent potential as an insect antifeedant.

TPS Genes Silencing Alters Constitutive Indirect and Direct Defense in Tomato

Following herbivore attacks, plants modify a blend of volatiles organic compounds (VOCs) released, resulting in the attraction of their antagonists. However, volatiles released constitutively may affect herbivores and natural enemies’ fitness too. In tomato there is still a lack of information on the genetic bases responsible for the constitutive release of VOC involved in direct and indirect defenses. Here we studied the constitutive emissions related to the two most abundant sesquiterpene synthase genes expressed in tomato and their functional role in plant defense. Using an RNA interference approach, we silenced the expression of TPS9 and TPS12 genes and assessed the effect of this transformation on herbivores and parasitoids. We found that silenced plants displayed a different constitutive volatiles emission from controls, resulting in reduced attractiveness for the aphid parasitoid Aphidius ervi and in an impaired development of Spodoptera exigua larvae. We discussed these data considering the transcriptional regulation of key-genes involved in the pathway of VOC metabolism. We provide several lines of evidence on the metabolic flux from terpenoids to phenylpropanoids. Our results shed more light on constitutive defenses mediated by plant volatiles and on the molecular mechanisms involved in their metabolic regulation.