Without roots there would be no rhizosphere and no rhizodeposition to fuel microbial activity. Although micro-organisms may view roots merely as a source of carbon supply this belies the fascinating complexity and diversity of root systems that occurs despite their common function. Here, we examine the physiological and genetic determinants of root growth and the complex, yet varied and flexible, root architecture that results. The main functions of root systems are also explored including how roots cope with nutrient acquisition from the heterogeneous soil environment and their ability to form mutualistic associations with key soil microorganisms (such as nitrogen fixing bacteria and mycorrhizal fungi) to aid them in their quest for nutrients. Finally, some key biotic and abiotic constraints on root development and function in the soil environment are examined and some of the adaptations roots have evolved to counter such stresses discussed.Keywords Root systems . Auxin . Root architecture . Soil heterogeneity . Abiotic and biotic stresses . Soil micro-organisms (including nitrogen-fixing bacteria and mycorrhizal fungi)Physiological and genetic determinants of root growth and achitecture A major difference between plant and animal development is that positional information rather than cell lineage determines cell fate in plants (Singh and Bhalla 2006). Post-embryonically, plant development is essentially driven by stem cells localized in apical regions of shoots and roots, and referred to as apical meristems. This particular characteristic allows plants, Plant Soil (