Primary Succession Shifts Fine-Root Nutrient Acquisition from Morphological Capture to Rhizosphere-Mediated Biochemical Mobilization

Primary succession following glacier retreat provides a natural system for testing whether soil development simply shifts fine roots along a single acquisitive–conservative axis orinstead changes the nutrient-acquisition pathway that dominates at the community level. We hypothesized a stage-dependent sequence, from substrate-limited exploration, to transient morphological capture, and finally to rhizosphere-mediated biochemical mobilization. To test this idea, we quantified fine-root morphology, absorptive-transport partitioning, anatomy, phosphatase activity, exudation, community-scale belowground structure, and soil and rhizosphere properties across woody communities representing approximately 20, 40, and 90 years since deglaciation in the Hailuogou Glacier foreland. Across succession stages, bulk density and pH declined, whereas field capacity, soil carbon, and soil nitrogen increased, indicating rapid development of the belowground resource environment. Fine-root strategies did not fall along a single acquisitive–conservative continuum. Instead, morphological nutrient capture peaked at intermediate succession: the 40-year stage had the highest specific root length, specific root area, absorptive-to-transport root length ratio, and root nitrogen concentration. In contrast, the 90-year stage showed lower specific root length but higher dry matter content, thicker cortex, greater standing fine-root biomass, larger rhizosphere volume, higher phosphatase activity, and greater area-based carbon exudation. This late-successional syndrome coincided with stronger extracellular enzyme activity, larger dissolved organic carbon and nitrogen pools, and higher microbial biomass, despite negative net nitrogen mineralization. Species-level analyses showed that biochemical-input traits were jointly shaped by successional stage, species identity, and their interaction. Together, these results show that primary succession did not simply increase or decrease root acquisitiveness. Instead, as soils developed, it changed the nutrient-acquisition pathway that dominated, with direct implications for nutrient cycling and vegetation dynamics in rapidly developing glacier-foreland ecosystems.