Vascular architecture and patchy nutrient availability generate within-plant heterogeneity in plant traits important to herbivores

Within-plant heterogeneity in growth, morphology, and chemistry is ubiquitous, and is commonly attributed to differences in tissue age, light availability, or previous damage by herbivores. Although these factors are important, we argue that plant vascular architecture is an underappreciated determinant of heterogeneity. Vascular architecture can restrict the transport of resources (nutrients, photosynthate, hormones, etc.) to within specific sectors of the plant: this is referred to as sectoriality. Although studies have documented sectoriality in the transport of isotopes and dyes from roots to shoots, the ecological consequences of this sectoriality remain poorly understood. We tested the hypothesis that spatial variation in belowground nutrient availability combined with sectorial transport results in localized "fertilization" of aboveground plant parts and generates heterogeneity in traits important to herbivores. Our split-root experiments with tomato (Lycopersicon esculentum Mill) clearly demonstrate that fertilization to isolated lateral roots generates heterogeneity in leaf morphology, phenolic chemistry, and side-shoot growth. Specifically, leaflets with direct connections to these lateral roots were larger and had lower levels of rutin and chlorogenic acid than did leaflets in other sectors lacking direct vascular connections. Moreover, side-shoot production was greater in the connected sectors. We discuss the implications of this heterogeneity for plant-herbivore interactions.