Link between defoliation and light treatments on root vitality of five understory shrubs with different resistance to insect herbivory

Understory shrubs are frequently attacked by insect herbivores. However, very little is known regarding possible interactions between light condition, defoliation (D) and fine root vitality (% live roots) and metabolic activity, and whether different plant strategies (compensation, trade-off and equilibrium) to defoliation depend on individual species light requirements. To explore the response of roots to such conditions, an experiment was established in which we experimentally removed 50% of leaves in 1-year-old seedlings of Sambucus nigra, Cornus sanguinea, Prunus serotina, Frangula alnus and Corylus avellana grown in 15% and full sunlight. On average, defoliation leads to a 15% reduction in fine root (<2 mm) vitality (% live roots). However, a statistically significant reduction in root vitality after defoliation was detected only in those species that are less herbivorized in nature (48% in S. nigra and 5% in C. sanguinea). On average, shade conditions (L) resulted in 18% decline in root vitality, and the effects of defoliation were also 22% higher than for plants grown in full light. Root vitality in both treatments (D and L) was significantly correlated with their dry mass, concentration of total phenol (TPh) and carbon to nitrogen ratio, and negatively correlated with nitrogen, soluble carbohydrates, starch and total non-structural carbohydrates (TNC). To a large extent, root vitality and chemistry varied by species. Higher root vitality was related to higher concentrations of phenolics, more than to N and TNC concentrations. Concentrations of phenolics also differed significantly between defoliated plants and controls. However, in defoliated plants, an increase in TPh was observed only in two species, which belong to two different groups in light requirements and susceptibility to insect grazing (C. sanguinea and P. serotina). This study indicated that higher vitality of roots occurred in species that are characterized by higher insect defoliation under natural conditions. It is likely that higher root vitality of these species was related to their high level of TPh and tannins. This was especially noticeable for the reduced light treatment, which represents natural conditions under which insect defoliation is highest. Our results suggest that varied strategies of resource allocation were used by the different species in response to variations in light and defoliation.