Context dependence in foraging behaviour of Achillea millefolium.

Context-dependent foraging behaviour is acknowledged and well documented for a diversity of animals and conditions. The contextual determinants of plant foraging behaviour, however, are poorly understood. Plant roots encounter patchy distributions of nutrients and soil fungi. Both of these features affect root form and function, but how they interact to affect foraging behaviour is unknown. We extend the use of the marginal value theorem to make predictions about the foraging behaviour of roots, and test our predictions by manipulating soil resource distribution and inoculation by soil fungi. We measured plant movement as both distance roots travelled and time taken to grow through nutrient patches of varied quality. To do this, we grew Achillea millefolium in the centers of modified pots with a high-nutrient patch and a low-nutrient patch on either side of the plant (heterogeneous) or patch-free conditions (homogeneous). Fungal inoculation, but not resource distribution, altered the time it took roots to reach nutrient patches. When in nutrient patches, root growth decreased relative to homogeneous soils. However, this change in foraging behaviour was not contingent upon patch quality or fungal inoculation. Root system breadth was larger in homogeneous than in heterogeneous soils, until measures were influenced by pot edges. Overall, we find that root foraging behaviour is modified by resource heterogeneity but not fungal inoculation. We find support for predictions of the marginal value theorem that organisms travel faster through low-quality than through high-quality environments, with the caveat that roots respond to nutrient patches per se rather than the quality of those patches.

Disentangling root system responses to neighbours: identification of novel root behavioural strategies.

Plants live in a social environment, with interactions among neighbours a ubiquitous aspect of life. Though many of these interactions occur in the soil, our understanding of how plants alter root growth and the patterns of soil occupancy in response to neighbours is limited. This is in contrast to a rich literature on the animal behavioural responses to changes in the social environment. For plants, root behavioural changes that alter soil occupancy patterns can influence neighbourhood size and the frequency or intensity of competition for soil resources; issues of fundamental importance to understanding coexistence and community assembly. Here we report a large comparative study in which individuals of 20 species were grown with and without each of two neighbour species. Through repeated root visualization and analyses, we quantified many putative root behaviours, including the extent to which each species altered aspects of root system growth (e.g. rooting breadth, root length, etc.) in response to neighbours. Across all species, there was no consistent behavioural response to neighbours (i.e. no general tendencies towards root over-proliferation nor avoidance). However, there was a substantial interspecific variation showing a continuum of behavioural variation among the 20 species. Multivariate analyses revealed two novel and predominant root behavioural strategies: (i) size-sensitivity, in which focal plants reduced their overall root system size in response to the presence of neighbours, and (ii) location-sensitivity, where focal plants adjusted the horizontal and vertical placement of their roots in response to neighbours. Of these, size-sensitivity represents the commonly assumed response to competitive encounters-reduced growth. However, location sensitivity is not accounted for in classic models and concepts of plant competition, though it is supported from recent work in plant behavioural ecology. We suggest that these different strategies could have important implications for the ability of a plant to persist in the face of strong competitors, and that location sensitivity may be a critical behavioural strategy promoting competitive tolerance and coexistence.