Heterogeneity of individual roots within the fine root architecture: causal links between physiological and ecosystem functions

This review covers the heterogeneity in functions within the fine root architecture in order to clarify the multiple functions of fine roots. Many fine root characteristics, such as anatomy, physiology, morphology, and their consequences for the ecosystem, differ among root ages and ontogenetic branching hierarchies. Individual root age can be characterized by tissue development, with the main tissues developing from primary to secondary tissues. The physiological characteristics of individual roots, such as absorptivity and respiration rates, decrease with increasing branching order, mainly because of aging and tissue development. The C/N ratio and lignin and suberin contents also increase with branching order because of root aging. Morphological characteristics, such as diameter and specific root length, differ among root orders because of both aging and ontogenetic differences. The mortality of individual roots differs among branching orders and root diameters. The life cycles of roots in the fine root architecture, that is, ephemeral and perennial, indicate ontogenetic differences in functions and demographic traits, similar to those for leaves and branches in shoots. In addition, differences in individual root life cycles may affect the root chemical composition, in turn, affecting the decomposition rate. Future studies should seek to identify heterorhizic units in mortality related to anatomical, physiological, and morphological differences for various species. The decomposition processes of each mortality unit within the fine root architecture are also important in understanding the link between physiological and ecosystem functions.