Fine root branch orders respond differentially to carbon source-sink manipulations in a longleaf pine forest

Abstract
Fine roots are a key component of carbon (C) flow and nitrogen (N) cycling in forest ecosystems. However, the complexity and heterogeneity of the fine root branching system have hampered the assessment and prediction of C and N dynamics at ecosystem scales. We examined how root morphology, biomass, and chemistry differed with root branch orders (1–5 with root tips classified as first order roots) and how different root orders responded to increased C sink strength (via N fertilization) and reduced carbon source strength (via canopy scorching) in a longleaf pine (Pinus palustris L.) ecosystem. With increasing root order, the diameter and length of individual roots increased, whereas the specific root length decreased. Total root biomass on an areal basis was similar among the first four orders but increased for the fifth order roots. Consequently, total root length and total root surface area decreased systematically with increasing root order. Fine root N and lignin concentrations decreased, while total non-structural carbohydrate (TNC) and cellulose concentrations increased with increasing root order. N addition and canopy disturbance did not alter root morphology, but they did influence root chemistry. N fertilization increased fine root N concentration and content per unit area in all five orders, while canopy scorching decreased root N concentration. Moreover, TNC concentration and content in fifth order roots were also reduced by canopy scorching. Our results indicate that the small, fragile, and more easily overlooked first and second order roots may be disproportionately important in ecosystem scale C and N fluxes due to their large proportions of fine root biomass, high N concentrations, relatively short lifespans, and potentially high decomposition rates.