“…Thus, branching density tends to decrease in the distal part of the root. The variation in branching density can also be attributed to soil factors, such as to gradients in soil bulk density (Konôpka et al, 2009), in soil moisture (Johnson and Aguirre, 1991;Ito et al, 2006), in soil temperature (Sattelmacher et al, 1990) and in soil nutrients (Gruber et al, 2013), including as roots traverse nutrient patches (Hodge, 2004;Yu et al, 2014). However, as these first-order LRs are located at different positions along the nodal roots, and the nodal roots from higher whorls exhibit steeper root angles, so that they penetrate the soil more vertically, the first-order LRs will experience a diversity of local soil environments, so being more likely to be affected by a range of soil factors.…”