In nature, essential resources for organisms, such as food for animals and light, water and nutrients for plants, are usually heterogeneously distributed, even at very small scale. As a result, all organisms, particularly plants mostly sessile, have a difficulty in acquiring essential resources from their environments. Animals express various types of foraging behavior to capture heterogeneously distributed essential foods. Clonal growth ( a vegetative reproductive process where by more than one individual of identical genetic composition is formed ) provides clonal plant not only with many “mouths” at different spatial positions, but also with a large spacial movability. As a clonal plant grows in environments characterized by a small scale resource heterogeneity, its inter ramet connection permits a resource sharing among the connected ramets. In addition, it may also allow certain ramets to respond locally and non locally to resousce heterogeneity. This may lead to a division of labor among the connected ramets and a selective placement of ramets in favorable micro habitats. Together these may enhance exploitation of resource heterogeneity by clonal plants, and in turn greatly contribute to maintenance or improvement of fitness. Such a behavior of clonal plants, expressed in heterogeneous environments, is to a large extent comparable to that of animals. Therefore, it has been considered as foraging behavior in clonal plants. More recently, it has been observed that phenotypic plasticity of clonal plants, which is relevant to foraging behavior, varies among species, types of genet architecture as well as among types of plants’ habitats. Foraging in clonal plants and its diversity have been receiving increasingly intensive investigations.
In a greenhouse experiment plasticity of clonal growth and clonal morphology of the stoloniferous rosette herb Halerpestes nahenica Ovcz. in response to differing levels of light intensity and nutrient availability was studied. Total plant dry weight, leaf area of primary ramets, total number of ramets and of stolons, and total stolon length were significantly reduced, while specific internode length and specific petiole length significantly increased under deep shading (6.25% of high light intensity, 5.3% of full daylight) or under low nutrient availability. Under low nutrient availability, mean stolon internode length of H. ruthenica was significantly larger while branching intensity and number of ramets smaller than those under high nutrient availability. These responses are consistent with the foraging model of clonal plants, indicating that H. nahenica is able to forage nutrients through the plastic responses of clonal growth and clonal. morphology when it grows in heterogeneous environments. However, under deep shading, both mean stolon internode length and mean petiole length were significantly reduced, which disagrees with the findings of many other stoloniferous herbs in response to low or medium levels of shading (ca. 13%-75% of high light intensity, >10% of full daylight), suggesting that under deep shading stoloniferous herbs may not forage light through the plastic responses of spacer length. Many traits such as total plant dry weight, total number and length of stolons, total length of secondary and tertiary stolons. total number of ramets, leaf area of primary ramets and branching intensity were markedly influenced by the interaction effect of light intensity and nutrient availability. Under high light intensity nutrient availability affected these traits more pronouncedly, however under low light intensity nutrient availability either did not affect or affected less markedly on these traits, indicating that fight intensity had significant effect on nutrient foraging in H. nahenica. Under deep shad
Environments with reciprocal patchiness of resources, in which the availability of two resources such as light and soil nutrients are patchily distributed in horizontal space and negatively correlated in each patch, are common in many ecosystems. The strategies by which clonal plants adapt to this type of heterogeneous environment were examined in three stoloniferous herbs,Potentilla reptans L. var. sericophylla Franch., P. anserina L. and Halerpestes ruthenica (Jacq.) Qvcz., commonly inhabiting forest understories, grasslands and low saline meadows, respectively. As pairs of connected ramets were subjected to reciprocal patchiness of light and nutrients, stolon connection between the two ramets significantly enhanced biomass of both ramet growing in low light intensity but high soil nutrient condition (LH ramet) and ramet growing in high light intensity but low soil nutrient condition (HL ramet) as well as whole ramet pairs (consisting of LH ramets and HL ramets). Additionally, stolon connection greatly increased root/shoot ratio of LH ramet while significantly decreased that of HL ramet. The results indicate that a reciprocal transportation of resources between interconnected ramets and a functional specialization of ramets in uptake of abundant resources occurred. By resource sharing and functional specialization, clonal plants can efficiently acquire locally abundant resources and buffer the stress caused by reciprocal patchiness of resources.
