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.

Responses of plants to herbivory are dependent on the type of damage and the ontogenetic stage of the plant. We compared the effects of stem pruning and defoliation on seedlings of Colophospermum mopane, an ecologically important tree species widely distributed in southern Africa. The growth of 160 greenhouse-grown juveniles were measured for 6-mo after germination and then 6-mo after treatments including 50% defoliation, 100% defoliation, 50% stem pruning and controls. Pruning resulted in 30% reductions in total leaf area, height and biomass. Partial defoliation resulted in 30% reductions in total leaf area and plant biomass. However, complete defoliation resulted in a 30% increase in biomass production, a doubling in leaf and lateral branch number, a 45% reduction in leaf size, and no change in total leaf area. Thus, completely defoliated seedlings showed greater performance than those that were only partially defoliated, indicating that C. mopane has become adapted to the chronic and severe defoliation inflicted by Imbrasia belina caterpillars. Comparison of our results with other studies indicates that C. mopane seedlings are less herbivory-tolerant than adults and that pruning has more negative effects than defoliation. Thus, seedling browsers may constrain recruitment in C. mopane, influencing its population dynamics and abundance.

Increased shoot growth has recurrently been recorded following real or simulated shoot browsing, although responses may vary between different woody species. We investigated morphological responses to simulated dry-season browsing on Colophospermum mopane, Acacia tortilis, Grewia flava, Peltophorum africanum and Ziziphus mucronata, which differ in growth rate, occurrence of spines and use by mammalian herbivores. Treatment groups included a control, four clipping intensities and cutting at ground level. Responses were monitored during the following growth season on three annual shoots, located in the bottom, middle and top height interval, on each sapling. All five species showed compensatory growth responses, although no obvious relation was found between the strength of responses and growth rate or natural browsing pressure. The general pattern was an increased shoot length, shoot diameter, number of lateral shoots and shoot biomass with increased clipping intensity. These responses are probably an effect of the reduction of competing growing points and disrupted apical dominance. Growth responses were stronger in shoots with higher location on the sapling, which may reflect the pressure on saplings to attain a certain height to avoid browsing, reproduce and increase resistance to fire. We found no evidence of the number of spines being an induced defence in either of the two spinescent species studied.

Field-grown trees of Alnus incana (L.) Moench, Alnus glutinosa (L.) Geartner and Betula pendula Roth displayed pronounced differences in responses of light-saturated net photosynthesis (Asat) to herbivory by the alder beetle (Agelastica alni L., Galerucinae), a specialized insect which primarily defoliates alders. We found that photosynthetic rates of grazed leaves increased following herbivory in Alnus but not in Betula. Area- and mass-based Asat of grazed leaves declined linearly with increasing amount of leaf perforation in B. pendula, by as much as 57%. By contrast Alnus glutinosa and Alnus incana increased area-based rates of Asat by 10–50% at all levels of leaf grazing. Given increased Asat in the remaining portion of grazed leaves, a net reduction in photosynthesis per leaf occurred only when the proportion of leaf area grazed exceeded 40% for Alnus incana and 23% for Alnus glutinosa. Since vein perforation by Agelastica alni was observed much more frequently in leaves of Betula than in Alnus, we hypothesized that declining Asat in herbivorized Betula was related to this disruption of water transport. A field experiment with artificial leaf perforation demonstrated a greater decline in Asat in vein-perforated Betula leaves than perforated leaves with midrib veins intact. However, regardless of leaf perforation regime, birch never showed post-perforation increases in Asat. In all species, rates of transpiration of grazed leaves linearly increased and water-use efficiency decreased with increased amount of leaf perforation. In grazed Alnus incana leaves, increasing leaf area consumption by Agelastica alni resulted in an increase of total phenols, a reduction in starch content and no changes in nitrogen concentration in the remaining portion. The increase in photosynthesis in Alnus incana might be related to declining leaf starch concentration or increasing stomatal conductance, but was unrelated to leaf nitrogen concentration. These gas exchange and leaf chemistry measurements suggest that in contrast to B. pendula, Alnus incana and Alnus glutinosa, which are the major host species for Agelastica alni, possess leaf-level physiological adaptations and defence mechanisms which can attenuate negative effects of herbivory by the alder leaf-beetle.