From rhizosphere to detritusphere - Soil structure formation driven by plant roots and the interactions with soil biota

Roots and the associated soil directly affected by root activity, termed the rhizosphere, have both been extensively studied and recognized for their crucial role in soil functioning. The formation of the rhizosphere is primarily driven by the effect of roots on shaping the physical structure of the soil, which in turn has direct feedbacks on the interactions between physical, biological and chemical processes. As a result, the rhizosphere is a hot spot for microbial activity, cycling of nutrients and turnover of organic matter. Despite the pivotal role of soil structure in controlling rhizosphere processes, we still lack a quantitative description and understanding of the interrelationships of root-systems and soil in the creation and stabilization of soil structure. We provide a comprehensive review of current knowledge and novel insights into processes that drive the formation and stabilization of soil structure in the rhizosphere. These processes are regulated by multiple indirect and direct pathways, involving root growth, the production of rhizodeposits and root hairs, as well as the activity of soil microorganisms and fauna. Further, we highlight that rhizosphere processes may persist and evolve after root death to an extent currently largely unknown. Finally, we identify five pertinent challenges that should be addressed to fully apprehend rhizosphere processes and thus harness the potential resilience of plant-soil interactions. These challenges include refining structural assessment and sampling of rhizosheaths, examining the rhizosphere in-situ and bridging the gap between solid phase and pore scale research. In our view, overcoming these obstacles can be accomplished by combining the power of imaging and isotopic approaches, especially at the field scale, encompassing diverse soils and plant species. The ultimate objective of future research should be to upscale rhizosphere processes by conducting more field experiments in concert with modeling efforts, under the umbrella of collaborative interdisciplinary research.