Slugs, willow seedlings and nutrient fertilization: intrinsic vigor inversely affects palatability

This study evaluates how preference by a generalist slug herbivore Arion subfuscus changes inversely with seedling size across three levels of fertilization for three full-sib families of willow seedlings. We analyzed seedlings for condensed tannin and protein concentration, and related these data to changes in palatability. In preference tests over time, leaf discs from more fertilized seedlings experienced an extended window of vulnerability compared to discs from less fertilized seedlings, which were also more tannin-rich. In a whole seedling selection study, slugs readily attacked smaller seedlings (B/5 cm) but rarely attacked taller seedlings ( /10 cm). However, a general difference in risk of damage close to 50% existed when comparing shorter and taller individuals within each family and level of fertilizer. The decrease in palatability with height of the seedlings was positively correlated with an increase in condensed tannin concentration. We found no effect of seedling size on protein concentration. Akaiki index criterion model comparisons suggested that only main effects were important for explaining seedling choice by slugs as well as the ratio between proteins and condensed tannins. Seedling size, had the largest effect, followed by fertilizer level and family. Surprisingly, seedling size and fertilizer treatment had opposite effects on palatability to slugs. Size decreased probability of damage, whereas fertilization extended the window of susceptibility. Because the seedlings were even-aged, differences in size are interpreted as differences in growth rate or vigor. The positive phenotypic correlation found between size and tannin production in the less preferred willow seedlings confirms that several plant defense traits may be selected for simultaneously, because fast growth may allow an early development of plant defenses. We discuss these results in the light of plant-defense theories that predict a negative correlation between the allocation to growth and the production of secondary defense compounds.