505I.506II.506III.508IV.512V.513VI.514515References515 Summary Fine roots acquire essential soil resources and mediate biogeochemical cycling in terrestrial ecosystems. Estimates of carbon and nutrient allocation to build and maintain these structures remain uncertain because of the challenges of consistently measuring and interpreting fine‐root systems. Traditionally, fine roots have been defined as all roots ≤ 2 mm in diameter, yet it is now recognized that this approach fails to capture the diversity of form and function observed among fine‐root orders. Here, we demonstrate how order‐based and functional classification frameworks improve our understanding of dynamic root processes in ecosystems dominated by perennial plants. In these frameworks, fine roots are either separated into individual root orders or functionally defined into a shorter‐lived absorptive pool and a longer‐lived transport fine‐root pool. Using these frameworks, we estimate that fine‐root production and turnover represent 22% of terrestrial net primary production globally – a c. 30% reduction from previous estimates assuming a single fine‐root pool. Future work developing tools to rapidly differentiate functional fine‐root classes, explicit incorporation of mycorrhizal fungi into fine‐root studies, and wider adoption of a two‐pool approach to model fine roots provide opportunities to better understand below‐ground processes in the terrestrial biosphere.

Tree growth resources and the efficiency of resource-use for biomass production determine the productivity of forest ecosystems. In nutrient-limited forests, nitrogen (N)-fertilization increases foliage [N], which may increase photosynthetic rates, leaf area index (L), and thus light interception (I C). The product of such changes is a higher gross primary production and higher net primary production (NPP). However, fertilization may also alter carbohydrate partitioning from below-to aboveground, increasing aboveground NPP (ANPP). We analyzed effects of long-term N-fertilization on NPP, and that of long-term carbon storing organs (NPP S) in a Pinus sylvestris forest on sandy soil, a wide-ranging forest type in the boreal region. We based our analyses on a combination of destructive harvesting, consecutive mensuration, and optical measurements of canopy openness. After eight-year fertilization with a total of 70 g N m-2 , ANPP was 27 ± 7% higher in the fertilized (F) relative to the reference (R) stand, but although L increased relative to its pre-fertilization values, I C was not greater than in R. On the seventh year after the treatment initiation, the increase of ANPP was matched by the decrease of belowground NPP (78 vs. 92 g C m-2 yr-1 ; ~17% of NPP) and, given the similarity of I C , suggests that the main effect of N-fertilization was changed carbon partitioning rather than increased canopy photosynthesis. Annual NPP S increased linearly with growing season temperature (T) in both treatments, with an upward shift of 70.2 g C m-2 yr-1 by fertilization, which also caused greater amount of unexplained variation (r 2 = 0.53 in R, 0.21 in F). Residuals of the NPP S-T relationship of F were related to growing season precipitation (P, r 2 = 0.48), indicating that T constrains productivity at this site regardless of fertility, while P is important in determining productivity where N-limitation is alleviated. We estimated that, in a growing season average T (11.5 ± 1.0 °C; 33-year-mean), NPP S response to N-fertilization will be nullified with P 31 mm less than the mean (325 ± 85 mm), and would double with P 109 mm greater than the mean. These results suggest that inter-annual H. Lim et al. 3 variation in climate, particularly in P, may help explaining the reported large variability in growth responses to fertilization of pine stands on sandy soils. Furthermore, forest management of long-rotation systems, such as those of boreal and northern temperate forests, must consider the efficiency of fertilization in terms of wood production in the context of changes in climate predicted for the region.

The tree root-mycorhizosphere plays a key role in resource uptake, but also in the adaptation of forests to changing environments. The adaptive foraging mechanisms of ectomycorrhizal (EcM) and fine roots of Picea abies, Pinus sylvestris and Betula pendula were evaluated along a gradient from temperate to subarctic boreal forest (38 sites between latitudes 48°N and 69°N) in Europe. Variables describing tree resource uptake structures and processes (absorptive fine root biomass and morphology, nitrogen (N) concentration in absorptive roots, extramatrical mycelium (EMM) biomass, community structure of root-associated EcM fungi, soil and rhizosphere bacteria) were used to analyse relationships between root system functional traits and climate, soil and stand characteristics. Absorptive fine root biomass per stand basal area increased significantly from temperate to boreal forests, coinciding with longer and thinner root tips with higher tissue density, smaller EMM biomass per root length and a shift in soil microbial community structure. The soil carbon (C) : N ratio was found to explain most of the variability in absorptive fine root and EMM biomass, root tissue density, N concentration and rhizosphere bacterial community structure. We suggest a concept of absorptive fine root foraging strategies involving both qualitative and quantitative changes in the root-mycorrhiza-bacteria continuum along climate and soil C : N gradients.

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