Studies with model plants such as Arabidopsis thaliana have revealed that phytohormones are central regulators of plant defense. The intricate network of phytohormone signaling pathways enables plants to activate appropriate and effective defense responses against pathogens as well as to balance defense and growth. The timing of the evolution of most phytohormone signaling pathways seems to coincide with the colonization of land, a likely requirement for plant adaptations to the more variable terrestrial environments, which included the presence of pathogens. In this review, we explore the evolution of defense hormone signaling networks by combining the model plant-based knowledge about molecular components mediating phytohormone signaling and cross talk with available genome information of other plant species. We highlight conserved hubs in hormone cross talk and discuss evolutionary advantages of defense hormone cross talk. Finally, we examine possibilities of engineering hormone cross talk for improvement of plant fitness and crop production.

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Bacterial wilt caused by the soil-borne pathogen Ralstonia solancearum is economically devastating, with no effective methods to fight the disease. This pathogen invades plants through their roots and colonizes their xylem, clogging the vasculature and causing rapid wilting. Key to preventing colonization are the early defense responses triggered in the host’s root upon infection, which remain mostly unknown. Here, we have taken advantage of a high-throughput in vitro infection system to screen natural variability associated to the root growth inhibition phenotype caused by R. solanacearum in Arabidopsis during the first hours of infection. To analyze the genetic determinants of this trait, we have performed a Genome-Wide Association Study, identifying allelic variation at several loci related to cytokinin metabolism, including genes responsible for biosynthesis and degradation of cytokinin. Further, our data clearly demonstrate that cytokinin signaling is induced early during the infection process and cytokinin contributes to immunity against R. solanacearum. This study highlights a new role of cytokinin in root immunity, paving the way for future research that will help understanding the mechanisms underpinning root defenses.

Although gibberellic acid (GA) is widely used in agriculture, it is unclear whether exogenous GA makes aphid-infested, mycorrhizal plants more susceptible to herbivory. This study investigates the role of GA in modulating defenses in barrel medic plants (Medicago truncatula) that are infested with pea aphids (Acyrthosiphon pisum) and colonized by the beneficial symbiont Rhizophagus intraradices. Mock- and R. intraradices-inoculated potted plants were grown in a topsoil: sand mix for 42 days and were treated with GA or solvent. Subsequently, plants were exposed to herbivory or no aphid herbivory for 36 h and 7 days. Afterwards, plant growth parameters, aphid fitness, and foliar phytohormone concentrations were measured. The results revealed that GA regulates plant defenses during arbuscular mycorrhizal (AM) fungus–plant–aphid interactions as aphids that fed for 7 days on mycorrhizal, GA-untreated plants weighed more than those that fed on mycorrhizal, GA-treated plants. No major differences were detected in phytohormone levels at 36 h. Overall, mycorrhizal plants showed more shoot biomass compared to non-mycorrhizal controls. The arbuscule density and fungal biomass of R. intraradices were not altered by exogenous GA and aphid herbivory based on molecular markers. This study indicates that exogenous GA may help reduce aphid fitness when feeding on mycorrhizal plants.

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