Soil nutrients are often heterogeneously distributed in space and time at scales relevant to individual plants, and plants can respond by selectively proliferating their roots within nutrient-rich patches. However, many environmental factors may increase or decrease the degree of root proliferation by plants. I explored how soil fertility, nitrogen (N) or phosphorus (P) limitation, and soil oxygen availability affected root system response to nutrient heterogeneity in forested wetland ecosystems of southeastern United States. Fine root biomass was not correlated with soil nutrient availability within wetland ecosystems, but was related to ecosystem-scale fertility. Root systems generally did not respond to P-rich patches in both floodplain (nutrient-rich) and depressional swamps (nutrient-poor) swamps, but results were inconclusive because the growth medium (sand) potentially hindered root growth. In floodplain forests, roots proliferated into N-rich patches but not P-rich patches, even though litterfall N:P ratios were > 15, which suggested that these ecosystems were P-limited. The combination of nutrient and oxygen heterogeneity affected root proliferation and biomass growth of three common floodplain forest species (Liquidambar styraciflua, Fraxinus pennsylvanica, and Nyssa aquatica) in a potted study, which was related to species' flood tolerance. My results suggest that the environmental context of plants can affect roots system response to nutrient heterogeneity in forested wetland ecosystems and highlights the need for field studies that investigate this phenomenon. Learning how environmental conditions affect plant response to nutrient heterogeneity at a fine-scale will provide better predictions of nutrient cycling, plant competition and succession, and forest productivity, which are important factors that determine carbon sequestration and timber production. / Ph. D.

Resources in the soil are heterogeneously distributed. We know that plant species differ in their root responses to nutrient patches and that these differences in foraging can influence plant competition. However, most studies of root-resource interactions overlook the potential top-down influence of root herbivores. While root herbivores can influence plant community structure, the extent to which they influence ecosystem-scale factors such as net primary production is unclear. In addition, little is known regarding root herbivore foraging behaviors and, more importantly, whether these foraging behaviors can actually influence species interactions. In this dissertation, I present a conceptual model of soil-root-herbivore interactions in which soil resource heterogeneity structures both root dynamics and the abundance and influence of root herbivores. I conducted two field and one greenhouse experiment examining this proposed model. The dissertation includes an introductory chapter (Chapter 1), a field study examining root responses to manipulations of soil fertility and root herbivory (Chapter 2), a greenhouse study that used plant species responses to heterogeneity to develop predictions about the role of root herbivores in mixed-species neighborhoods (Chapter 3), and a field study of planted communities examining soil fertility and fauna effects on above- and belowground structure and function (Chapter 4). In all cases, there were significant effects of root herbivores on community structure and components of net primary production. Resource distribution had a strong effect in studies conducted in sandy, nutrient-poor soils (Chapter 2 and 3), but had a reduced effect in the study conducted at Kentland Farm in loamy soils (Chapter 4). Interactions between resource availability and root herbivory were common. These results support the theory that the potential benefit of resource-rich patches may be constrained by root herbivores. This research complements recent findings that demonstrate other potential costs of species foraging behaviors (such as exposure to soil anoxia and increased drought stress), as well as potential effects of root herbivores and other soil fauna on plant diversity. / Ph. D.

Root exploitation of enriched soil microsites was examined for the tussock grasses Agropyron desertorum and Agropyron spicatum and the shrub Artemisia tridentata. Two mechanisms of exploitation of the microsites were examined: root proliferation and changes in nutrient uptake capacity. One day after nutrient solution was applied to small soil patches, the mean relative growth rate of Agropyron desertorum roots in enriched patches was two to four times greater than for roots of the same plants in soil patches treated with distilled water. This rapid and striking root proliferation occurred in response to N-P-K enrichment as well as to P or N enrichment alone. Agropyron spicatum showed no tendency to proliferate roots in enriched soil patches during the two-week experiments. The shrub Artemisia tridentata proliferated roots within one day of initial solution injection in the N-enrichment experiment, but root proliferation of this species was more gradual and less consistent in other experiments. The ability of Agropyron desertorum partly explain its to proliferate superior ability roots to rapidly may exploit soil nutrients compared to Agropyron spicatum in Great Basin rangelands of North America. Changes in nutrient uptake capacity in enriched soil patches were also studied for each species. rapid changes in uptake capacity of plant roots Large and from the field were observed after creation of nutrient-rich patches in the soil. Phosphate uptake of excised roots from enriched soil patches was roots of control patches as much as 80% greater than for treated with distilled water. These increases in uptake capacity took place within one week of patch treatment for all three species. A follow-up experiment showed increases within three days of patch treatment. These results showing rapid physiological plasticity in roots exploiting nutrient patches have important implications for nutrient belowground competition among plants.