Plant root system response to nutrient heterogeneity was tested in ten plant species of varying life form and successional status. All plants tested are native to the South Carolina coastal plain. Morphological responses of the root system (scale, precision and discrimination) and overall plant response (sensitivity) to increasing nutrient heterogeneity were tested. Ten individuals of each species were placed into four treatments which had varying nutrient distribution but the same overall nutrient addition. Plants were harvested when roots reached pot edge. I observed high variation in scale (mass and extent of a root system), precision (the ability to proliferate roots in nutrient patches) and sensitivity (growth benefits gained as nutrient heterogeneity increases; measured as total biomass). No significant discrimination responses were observed, although greatest mean root density occurred at intermediate fertility levels for all species. I tested the hypothesis that scale and precision would be negatively correlated, and I did not observe this relationship in these plant species. However, in herbaceous species scale and precision were positively correlated. Sensitivity was not closely related to precision indicating that proliferating roots in fertile patches does not always yield growth benefits in heterogeneous soils. Further, some sensitive species had very low precision suggesting that other characteristics lead to positive growth response in heterogeneous environments. Plasticity of root uptake rates and demography of roots are proposed as two other mechanisms which may play important roles in plant sensitivity responses. Scale was negatively correlated to sensitivity for herbaceous plants suggesting that plants that monopolize the most soil space are not able to gain benefits from nutrient patches within the soil matrix. There was no trend observed to suggest that plant life form was correlated with precision or sensitivity. However, scale was greater in herbs than in woody plants, possibly because the two life forms develop at different times. / Master of Science

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.

Relationships between nutrient heterogeneity, root foraging behavior and short-term competitive interactions were investigated for six species native to southeastern USA. Monoculture, two- and six-species garden plots were established and fertilized to create spatially homogeneous or heterogeneous nutrient conditions. After 3.5 months, root proliferation in rich patches (precision) and aboveground biomass response to heterogeneity were assessed in monocultures, and competitive outcomes (aboveground biomass) were determined from mixed-species plots. In monoculture plots, two species were relatively precise foragers, but no species showed significant aboveground biomass response to nutrient treatment. Correlations between precision and aboveground biomass were weak (-0.40 < r < 0.17). In two-species plots, interspecific competition was influenced by soil heterogeneity in two of six cases tested (P < 0.05), and precision was the behavior most correlated with competitive success. In six-species plots, spatial pattern of nutrients had no influence on aboveground growth or competition. Results suggest that heterogeneity influences competition, but the influence is context-specific and generally small. Precision may be the foraging behavior that most influences interspecific interactions. / Ph. D.