Abstract
Plant-soil interactions have bottom-up and top-down effects within a plant community. Heavy metal pollution can change plant-soil interactions, directly influence bottom-up effects and indirectly affect herbivores within the community. In turn, herbivores can affect plant-soil interactions through top-down effects. However, the combined effects of heavy metals and herbivores on soil enzymes, plants and herbivores have rarely been reported. Therefore, the effects of lead (Pb), Spodoptera litura and their combined effects on soil enzyme activities, pakchoi nutrition, defence compounds and S. litura fitness were examined here. Results showed that Pb, S. litura and their combined effects significantly affected soil enzymes, pakchoi and S. litura. Specifically, exposure to double stress (Pb and S. litura) decreased soil urease, phosphatase and sucrase activities compared with controls. Furthermore, the soluble protein and sugar contents of pakchoi decreased, and the trypsin inhibitor content and antioxidant enzyme activity increased. Finally, the S. litura development period was extended, and survival, emergence rates and body weight decreased after exposure to double stress. The combined stress of Pb and S. litura significantly decreased soil enzyme activities. Heavy metal accumulation in plants may create a superposition or synergistic effect with heavy metal-mediated plant chemical defence, further suppressing herbivore development. Pb, S. litura and their combined effects inhibited soil enzyme activities, improved pakchoi resistance and reduced S. litura development. The results reveal details of soil-plant-herbivore interactions and provide a reference for crop pest control management in the presence of heavy metal pollution.
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