[Response of fine roots to soil nutrient spatial heterogeneity]

The spatial heterogeneity is the complexity and variation of systems or their attributes, and the heterogeneity of soil nutrients is ubiquitous in all natural ecosystems. The scale of spatial heterogeneity varies considerably among different ecosystems, from tens of centimeters to hundred meters. Some of the scales can be detected by individual plant. Because the growth of individual plants can be strongly influenced by soil heterogeneity, it follows that the inter-specific competition should also be affected. During the long process of evolution, plants developed various plastic responses with their root system, including morphological, physiological and mycorrhizal plasticity, to maximize the nutrient acquisition from heterogeneous soil resources. Morphological plasticity, an adjustment in root system spatial allocation and architecture in response to spatial heterogeneous distribution of available soil resources, has been most intensively studied, and root proliferation in nutrient rich patches has been certified for many species. The species that do respond may have an increased rate of nutrient uptake, leading to a competitive advantage. Scale and precision are two important features employed in describing the size and foraging behavior of root system. It was hypothesized that scale and precision is negatively related, i. e., the species with high scale of root system tend to be a less precise forager. The outcomes of different research work have been diverse, far from reaching a consensus. Species with high scale are not necessarily less precise in fine root allocation, and vice versa. The proliferation of fine root in enriched micro-sites is species dependent, and also affected by other factors, such as patch attributes (size and nutrients concentration), nutrients, and overall soil fertility. Beside root proliferation in nutrient enriched patches, plants can also adapt themselves to the heterogeneous soil environment by altering other root characteristics such as fine root diameter, branch angle, length, and spatial architecture of root system. Physiological and mycorrhizal plasticity can add some influence on the morphological plasticity to some extent, but they are less studied. Roots located in different patches can quickly regulate their nutrient uptake kinetics within different nutrient patches, and increase overall nutrient uptake. Physiological response may, to certain extent, reduce morphological response, and is meaningful for plant growth on soils with frequently changing spatial and temporal heterogeneity. Mycorrhizal plasticity has been least studied so far. Some researches revealed that mycorrhiza, rather than fine root, proliferated in enriched patches. But, it is not the case with other studies. The proliferation of mycorrhiza within enriched patches is more profitable in term of carbon invest. The effect of fine root proliferation on nutrient uptake is complex, depending on ion mobility and whether or not neighboring plant exists. The influence of root plasticity on the growth of plants is species specific. Some species (sensitive species) gain growth benefit, while others don't. The ability of an individual plant to response to heterogeneous resources has significant effect on its competitive ability and its fate within the community, and eventually shapes the composition and structure of the community.