Moderate drought alters biomass and depth distribution of fine roots in Norway spruce

A rain shelter experiment was conducted in a 90‐year‐old Norway spruce stand, in the Kysucké Beskydy Mts (Slovakia). Three rain shelters were constructed in the stand to prevent the rainfall from reaching the soil and to reduce water availability in the rhizosphere. Fine root biomass and necromass were repeatedly measured throughout a growing season by soil coring. We established the quantities of fine root biomass (live) and necromass (dead) at soil depths of 0–5, 5–15, 15–25 and 25–35 cm. Significant differences in soil moisture contents between control and drought plots were found in the top 15 cm of soil after 20 weeks of rainfall manipulation (lasting from early June to late October). Our observations show that even relatively light drought decreased total fine root biomass from 272.0 to 242.8 g m^?2 and increased the amount of necromass from 79.2 to 101.2 g m^?2 in the top 35 cm of soil. Very fine roots (VFR), that is, those with diameter up to 1 mm, were more affected than total fine roots defined as 0–2 mm. The effect of reduced water availability was depth‐specific; as a result, we observed a modification of vertical distribution of fine roots. More roots in drought treatment were produced in the wetter soil horizons at 25–35 cm depth than at the surface. We conclude that fine and VFR systems of Norway spruce have the capacity to re‐allocate resources to roots at different depths in response to environmental signals, resulting in changes in necromass to biomass ratio.     

Stand responses to initial spacing in Norway spruce plantations in Norway

Abstract

Using data from nine spacing experiments of Norway spruce (Picea abies (L.) Karst.) in Norway, covering wide ranges of site index and initial spacing, this study evaluated stand basal area and volume responses to initial spacing and examined whether these responses varied by stand ages or site quality. We developed nonlinear regression models that described the standing basal area or volume responses to initial spacing along with site index and stand ages. The results show that closer spacings produced higher standing basal area and volume than wider spacings. The response curves are highly nonlinear in younger stands and become nearly linear in mid-rotation stands, indicating stronger responses at younger ages and weaker responses as age increases. Furthermore, for young stands, spacing effects are stronger at closer than at wider spacings. The basal area and volume responses to initial spacing tend to be similar across site indices. However, the interaction of site index and stand age on spacing responses makes it difficult to isolate the effect of site index on spacing responses. Mortality is higher and begins earlier at closer spacings than wider. The mean diameter of the largest 100, 400, 600, and 800 trees ha^?1 increased with spacing in three out of the nine experiments. Dominant height did not vary by initial spacing for any of the experiments. The findings suggest that the extra volume production in stands of closer initial spacing is restricted to early stand development.     

Effects of slash retention and wood ash addition on fine root biomass and production and fungal mycelium in a Norway spruce stand in SW Sweden

In the study reported here we examined the short-term effects (1–3 years) of slash retention (SR) and the long-term effects (13–15 years) of wood-ash application (A) on fine roots and mycorrhizae in a 40-year-old Norway spruce forest in southwest Sweden. Soil cores were used to obtain estimates of the biomass (g m⁻²) of roots in three diameter classes (<0.5, 0.5–1 and 1–2 mm), root length density (RLD), specific root length (SRL) and mycorrhizal root tip density (RTD). Fine root (<1 mm) length production and mortality, and mycelium production, were estimated using minirhizotron and mesh bag techniques, respectively. Compared with the control plots (C), the biomass of fine roots in diameter classes <0.5 mm and 0.5–1 mm was significantly higher in A plots, but lower in SR plots. In addition, RLD was significantly lower in the humus layer of SR plots than in the humus layers of C and A plots, but not in the other layers. None of the treatments affected the SRL. In all soil layers, the SR treatment resulted in significant reductions in the number of ectomycorrhizal root tips, and the mycelia production of fungi in mesh bags, relative to the C treatment, but the C and A treatments induced no significant changes in these variables. Fine root length production in the C, A and SR plots amounted to 94, 87 and 70 mm tube⁻¹ during the 2003 growing season, respectively. Fine root mortality in treated plots did not change over the course of the study. We suggest that leaving logging residues on fertile sites may result in nitrogen mineralisation, which may in turn induce reductions in root biomass, and both root and mycelium production, and consequently affect nutrient uptake and the accumulation of organic carbon in soil derived from roots and mycorrhizae.     

Effects of experimental drought on the fine root system of mature Norway spruce

Norway spruce (Picea abies (L.) Karst.) is an economically important, but relatively drought-sensitive tree species that might suffer from increasing drought intensities and frequencies, which are predicted to occur in parts of central Europe under future climatic change. In a throughfall exclusion experiment using sub-canopy roofs, we tested the hypothesis that enhanced drought leads to an increase in fine root mortality, and also to a higher, subsequent fine root growth. Fine root production and mortality were assessed using two independent approaches, sequential soil coring (organic layer) and direct observations in minirhizotrons (organic layer plus upper mineral soil). Six weeks of throughfall exclusion resulted in mild drought stress, i.e. a reduction in average soil moisture from 20 to 12 vol.% during the treatment. Based on the sequential coring data, experimental drought did not result in significant changes in fine root biomass during the 6-week treatment period, but caused an increase in fine root mortality by 61% in the 6 weeks following the drought treatment. Remarkably, fine root production showed a synchronous increase in this period, which more than compensated for the loss due to increased mortality. The minirhizotron data confirmed that the drought treatment increased fine root loss in the organic layer. Based on this method, however, root loss occurred during the drought period and was not compensated by increased root production. The mild drought stress was mainly restricted to the organic layer and did not significantly influence fine root dynamics in the mineral soil. We calculated that the drought event resulted in an extra input of about 28 g C m⁻² and 1.1 g N m⁻² to the soil due to increased fine root mortality. We conclude that even periods of mild drought significantly increase fine root mortality and the associated input of root-derived C to the soil organic matter pool in temperate Norway spruce forests.     

Fine root biomass in relation to site and stand characteristics in Norway spruce and Scots pine stands

Variations in fine root biomass of trees and understory in 16 stands throughout Finland were examined and relationships to site and stand characteristics determined. Norway spruce fine root biomass varied between 184 and 370 g m(-2), and that of Scots pine ranged between 149 and 386 g m(-2). In northern Finland, understory roots and rhizomes (< 2 mm diameter) accounted for up to 50% of the stand total fine root biomass. Therefore, the fine root biomass of trees plus understory was larger in northern Finland in stands of both tree species, resulting in a negative relationship between fine root biomass and the temperature sum and a positive relationship between fine root biomass and the carbon:nitrogen ratio of the soil organic layer. The foliage:fine root ratio varied between 2.1 and 6.4 for Norway spruce and between 0.8 and 2.2 for Scots pine. The ratio decreased for both Norway spruce and Scots pine from south to north, as well as from fertile to more infertile site types. The foliage:fine root ratio of Norway spruce was related to basal area and stem surface area. The strong positive correlations of these three parameters with fine root nitrogen concentration implies that more fine roots are needed to maintain a certain amount of foliage when nutrient availability is low. No significant relationships were found between stand parameters and fine root biomass at the stand level, but the relationships considerably improved when both fine root biomass and stand parameters were calculated for the mean tree in the stand. When the northern and southern sites were analyzed separately, fine root biomass per tree of both species was significantly correlated with basal area and stem surface area per tree. Basal area, stem surface area and stand density can be estimated accurately and easily. Thus, our results may have value in predicting fine root biomass at the tree and stand level in boreal Norway spruce and Scots pine forests.     

Changes in fine root production and longevity in relation to water and nutrient availability in a Norway spruce stand in northern Sweden

Effects of irrigation and liquid fertilization on fine root (< 1 mm) production and longevity, and fine root (< 0.5-2 mm) biomass were studied in a Norway spruce (Picea abies (L.) Karst.) stand in northern Sweden. Fine root length production and longevity were measured by the minirhizotron technique at 0-10 cm depth in the following treatments: irrigation (I), liquid fertilization (IL) and control (C). Standing root biomass and root length density (RLD) were studied in the litter-fermented humus (LFH) layer and at depths of 0-10, 10-20 and 20-30 cm using soil cores in solid fertilized (F) and C plots. Minirhizotrons were installed in October 1994 and measurements recorded monthly from July to September 1995 and during the growing season in 1996. Soil cores were sampled in 1996. Fine root production increased significantly in IL plots compared with C plots, but the I treatment did not increase root production. Root mortality increased significantly in IL plots compared with C plots. Fine root longevity in IL plots was significantly lower compared with C and I plots. No significant difference was found between longevity of fine roots in I and C plots. Compared with C, F treatment increased fine root biomass in the LFH and mineral soil layers, and increased the amount of fine roots in mineral soil layers relative to the LFH layer. Furthermore, F increased RLD and the number of mycorrhizal root tips significantly.     

Changes in fine root production and longevity in relation to water and nutrient availability in a Norway spruce stand in northern Sweden

The PEACH computer simulation model of reproductive and vegetative growth of peach trees (Grossman and DeJong 1994) was adapted to estimate seasonal nitrogen (N) dynamics in organs of mature peach (Prunus persica (L.) Batsch cv. O'Henry) trees grown with high and low soil N availability. Seasonal N accumulation patterns of fruits, leaves, stems, branches, trunk and roots of mature, cropping peach trees were modeled by combining model predictions of organ dry mass accumulation from the PEACH model with measured seasonal organ N concentrations of trees that had been fertilized with either zero or 200 kg N ha(-1) in April. The results provided a comparison of the N use of perennial and annual organs during the growing season for trees growing under both low and high N availability. Nitrogen fertilization increased tree N content by increasing organ dry masses and N concentrations during the fruit growing season. Dry mass of current-year vegetative growth was most affected by N fertilization. Whole-tree N content of fertilized trees was almost twice that of non-fertilized trees. Although N use was higher in fertilized trees, calculated seasonal N accumulation patterns were similar for trees in both treatments. Annual organs exhibited greater responses to N fertilization than perennial organs. Estimated mean daily N use per tree remained nearly constant from 40 days after anthesis to harvest. The calculations indicated that fertilized trees accumulated about 1 g N tree(-1) day(-1), twice that accumulated by non-fertilized trees. Daily N use by the fertilized orchard was calculated to be approximately 1 kg N ha(-1), whereas it was approximately 0.5 kg N ha(-1) for the non-fertilized trees. During the first 25-30 days of the growing season, all N use by growing tissues was apparently supplied by storage organs. Nitrogen release from storage organs for current growth continued until about 75 days after anthesis in both N treatments.     

Comparing growth and fine root distribution in monocultures and mixed plantations of hybrid poplar and spruce

Disease prevention, biodiversity, productivity improvement and ecological considerations are all factors that contribute to increasing interest in mixed plantations. The objective of this study was to evaluate early growth and productivity of two hybrid poplar clones, P. balsamifera x trichocarpa (PBT) and P. maximowiczii x balsamifera (PMB), one improved family of Norway spruce (Picea glauca (PA)) and one improved family of white spruce (Picea abies (PG)) growing under different spacings in monocultures and mixed plots. The plantations were established in 2003 in Abitibi-Témiscamingue, Quebec, Canada, in a split plot design with spacing as the whole plot factor (1 × 1 m, 3 × 3 m and 5 × 5 m) and mixture treatments as subplot factor (pure: PBT, PMB, PA and PG, and 1:1 mixture PBT:PA, PBT:PG, PMB:PA and PMB:PG). Results showed a beneficial effect of the hybrid poplar-spruce mixture on diameter growth for hybrid poplar clones, but not for the 5 × 5 m spacing because of the relatively young age of the plantations. Diameter growth of the spruces decreased in mixed plantings in the 1 × 1 m, while their height growth increased, resulting in similar aboveground biomass per tree across treatments. Because of the large size differences between spruces and poplars, aboveground biomass in the mixed plantings was generally less than that in pure poplar plots. Leaf nitrogen concentration for the two spruce families and hybrid poplar clone PMB was greater in mixed plots than in monocultures, while leaf nitrogen concentration of clone PBT was similar among mixture treatments. Because of its faster growth rate and greater soil resources demands, clone PMB was the only one showing an increase in leaf N with increased spacing between trees. Fine roots density was greater for both hybrid poplars than spruces. The vertical distribution of fine roots was insensitive to mixture treatment.     

Tree age effect on fine-root and leaf morphology in a silver birch forest chronosequence

The influence of forest ageing on fine-root morphology and relations between fine-root and leaf characteristics is poorly studied. The aim of this study was to analyse age-driven changes in ectomycorrhizal roots (EcM roots) and leaf morphology in a chronosequence of silver birch (Betula pendula Roth.), which would provide a better understanding of adaptation responses and acclimation capacity of tree roots and leaves. The chronosequence included six age classes (3, 6, 14, 32, 45, and 60 years.). All stands had regenerated naturally and grew in a highly productive Oxalis forest site type in Estonia. Most changes in the morphology of EcM roots and leaves of silver birch occur faster at a young age. The functional parameters—mean specific area of EcM root (SRA) and leaf specific area (SLA) as well as leaf N—decreased with age. EcM root SRA and specific root length (SRL) decreased with stand age as a result of increased mean diameter and tissue density. In age classes of 6, 14, and 32 years, the total number of dominating EcM taxa was 34, and the distribution of four different dominating EcM exploration types (contact-, short-, medium-, long-distance) was similar. We conclude that high values of SRA, SLA, and leaf N measured in young silver birch stands indicate high activity of physiological processes necessary for fast-growing young trees. A decrease of SLA and SRA and N in the chronosequence of fertile stands of silver birch is most probably caused by down-regulation of growth, affecting simultaneously leaves and fine roots.     

Contribution of root respiration to soil surface CO2 flux in a boreal black spruce chronosequence

We quantified the contributions of root respiration (RC) and heterotrophic respiration to soil surface CO2 flux (RS) by comparing trenched and untrenched plots in well-drained and poorly drained stands of a black spruce (Picea mariana (Mill.) BSP) fire chronosequence in northern Manitoba, Canada. Our objectives were to: (1) test different equations for modeling RS as a function of soil temperature; and (2) model annual RS and RC for the chronosequence from continuous soil temperature measurements. The choice of equation to model RS strongly affected annual RS and RC, with an Arrhenius-based model giving the best fit to the data, especially at low temperatures. Modeled values of annual RS were positively correlated with soil temperature at 2-cm depth and were affected by year of burn and trenching, but not by soil drainage. During the growing season, measured RC was low in May, peaked in late July and declined to low values by the end of the growing season. Annual RC was < 5% of RS in the recently burned stands, approximately 40% in the 21-year-old stands and 5-15% in the oldest (152-year-old) stands. Evidence suggests that RC may have been underestimated in the oldest stands, with residual root decay from trenching accounting for 5-10% of trenched plot RS at most sites.     

Quantification of two-year-old hybrid poplar root systems: morphology, biomass, and (14)C distribution

Root morphology, biomass, and (14)C distribution were studied in two 2-year-old Populus trichocarpa x P. deltoides hybrids, which originated from hardwood cuttings, to determine the pattern of root distribution in a plantation and to refine methods for root recovery. The trees were labeled with (14)CO(2) and harvested after a 72-hour chase period. Roots attached to each labeled tree were analyzed for morphological traits at the time of harvest. Detached roots from within a 1-m(3) volume of soil surrounding each tree were separated from the soil and sorted on the basis of rooting depth and root diameter. Lateral roots > 2 mm in diameter had a largely horizontal orientation at their point of origin from the cutting and extended horizontally up to 4 m from the cutting. This resulted in considerable overlap of root systems in the plantation. Results from (14)C labeling indicated that 24 +/- 4% (+/- SD) of the carbon exported from branches-labeled within two weeks after branch budset-was translocated to the root system. Dilution of the root (14)C label indicated that from 0 (> 5 mm diameter roots) to 75% (< 2 mm diameter roots) of the roots recovered from within the 1-m(3) volume of soil surrounding a harvested tree originated from other trees. Total root biomass was 6 +/- 1 Mg ha(-1) for both hybrids. Sixty percent of the root biomass was recovered directly from excavation, 16% from coarse-sieving excavated soil, and 24% from re-sorting sieved soil. The study indicated that root growth of hybrid poplars may be rapid and extensive and that detailed sorting of soil subsamples substantially improves the recovery of fine roots < 2 mm in diameter.     

Inherited narrow crown form, harvest index and stem biomass production in Norway spruce, Picea abies

The effect of crown form on stem biomass production was investigated in an 18-19-year-old Norway spruce stand (Picea abies (L.) Karst.). The harvest index was 0.271 in pendula trees, which have a heritable narrow crown form, and 0.235 in normal-crowned trees and the dry weights of stem biomass were 2.57 kg and 3.37 kg, respectively. However, the production of stem biomass per crown projected area was more than twice as much in pendula trees as in normal-crowned trees. Results indicate that the crown form of pendula spruce is less plastic than that of normal-crowned spruce suggesting that such trees might maintain a high stemwood yield per unit ground area with increasing stand density. Because pendula spruce has a higher harvest index and a less flexible crown than normal-crowned spruce, it might be useful for crop tree ideotype breeding.     

Variation patterns of fine root biomass,production and turnover in Chinese forests

China’s forests cover 208.3 million ha and span a wide range of climates and a large variety of forest types including tropical,temperate,and boreal forests.However the variation patterns of fine root(2 mm in diameter biomass,production,and turnover from the south to the north are unclear.This study summarizes fine root biomass(FRB),production(FRP)and turnover rate(FRT)in Chi na’s forests as reported by 140 case studies published from 1983 to 2014.The results showed that the mean values o FRB,FRP and FRT in China’s forests were 278 gm~(-2)366 gm~(-2)a~(-1),and 1.19 a~(-1),respectively.Compared with other studies at the regional or global scales,FRB in China’s forests was lower,FRP was similar to estimates a the global scale,but FRT was much higher.FRB,FRP,and FRT in China’s forests increased with increasing mean annual precipitation(MAP),indicating that fine root vari ables were likely related to MAP,rather than mean annua temperature or latitude.This is possibly due to the smal variation in temperature but greater variation in precipitation during the growing season.These findings suggest that spatiotemporal variation in precipitation has a more profound impact on fine root dynamics in China’s forests,and this will impact carbon and nutrient cycles driven by root turnover in the future.     

Spatial distribution and persistence of Heterobasidion parviporum genets on a Norway spruce site

Spatial distribution of Heterobasidion genets over a period of ca 50 years in two successive generations of Norway spruce (Picea abies) was unravelled. The genets were first identified in 1993 in a naturally regenerated 43‐year‐old spruce stand that had been thinned the previous winter. Heterobasidion parviporum was found in 17.5% of the old stumps of the previous spruce generation. Nine genets were identified on the study plot; seven of them were present in old stumps of the previous tree generation and two only in the new spruce generation. Eighteen spruce trees of the new generation were infected, 15 of them by vegetative growth of genets originating from the old stumps. The study plot was investigated again in 2005. No new genets had been established after thinning, and three old genets had died out. The remaining genets had infected five new trees, most likely from the thinning stumps of diseased trees. At the age of 56 years, 16.1% of the residual spruces were infected by Heterobasidion. The results of this study suggest that if spore infection to stumps of spruce can be prevented, the decay frequency caused by H. parviporum will not necessarily increase in successive generations.     

Fine root biomass,distribution, and production in young pine-hardwood stands

Patterns of fine root biomass, production, and distribution were estimated for pure stands and mixtures of three-year-old loblolly pine (Pinus taeda L.) with red maple (Acer rubrum L.) or black locust (Robinia pseudoacacia L.) on the Virginia Piedmont to determine the role of fine roots in interference between pine and hardwood tree species. Estimates were based on amounts of live and dead fine roots separated from monthly core samples during the third growing season after planting. Live and dead fine root biomass and production varied by species, but mixtures of loblolly pine and black locust generally had greater fine root biomass and fine root production than pure stands or loblolly pine-red maple mixtures. Hardwood species had greater live fine root biomass per tree in mixtures with pine compared to pure stands. Greater live fine root biomass in pine-locust stands may be attributed to differential utilization of the soil volume by fine roots of these species. For all stands, approximately 50% of live five root biomass was located in the upper 10 cm of soil.     

Fine root dynamics, coarse root biomass, root distribution, and soil respiration in a multispecies riparian buffer in Central Iowa, USA

By influencing belowground processes, streamside vegetation affects soil processes important to surface water quality. We conducted this study to compare root distributions and dynamics, and total soil respiration among six sites comprising an agricultural buffer system: poplar (Populus × euroamericana‘ Eugenei), switchgrass, cool-season pasture grasses, corn (Zea mays L.), and soybean (Glycine max (L.) Merr.). The dynamics of fine (0--2 mm) and small roots (2--5 mm) were assessed by sequentially collecting 35 cm deep, 5.4 cm diameter cores from April through November. Coarse roots were described by excavating 1 × 1 × 2 m pits and collecting all roots in 20 cm depth increments. Root distributions within the soil profile were determined by counting roots that intersected the walls of the excavated pits. Soil respiration was measured monthly from July to October using the soda-lime technique. Over the sampling period, live fine-root biomass in the top 35 cm of soil averaged over 6 Mg ha^-1 for the cool-season grass, poplar, and switchgrass sites while root biomass in the crop fields was < 2.3 Mg ha^-1 at its maximum. Roots of trees, cool-season grasses, and switchgrass extended to more than 1.5 m in depth, with switchgrass roots being more widely distributed in deeper horizons. Root density was significantly greater under switchgrass and cool-season grasses than under corn or soybean. Soil respiration rates, which ranged from 1.4--7.2 g C m^-2 day^-1, were up to twice as high under the poplar, switchgrass and cool-season grasses as in the cropped fields. Abundant fine roots, deep rooting depths, and high soil respiration rates in the multispecies riparian buffer zones suggest that these buffer systems added more organic matter to the soil profile, and therefore provided better conditions for nutrient sequestration within the riparian buffers. This revised version was published online in June 2006 with corrections to the Cover Date.     

Factors causing variation in fine root biomass in forest ecosystems

Fine roots form one of the most significant components contributing to carbon cycling in forest ecosystems. We study here the effect of variation in root diameter classes, sampling depth and the inclusion of understorey vegetation root biomass in fine root biomass (FRB) estimates. The FRB estimates for different forest biomes are updated using a database of 512 forest stands compiled from the literature. We also investigate the relationships between environmental or forest stand variables and fine root biomass (≤2 mm in diameter) at the stand (g m⁻²) and tree level (g tree⁻¹). The FRB estimates extrapolated for the whole rooting depth were 526 ± 321 g m⁻², 775 ± 474 g m⁻² and 776 ± 518 g m⁻² for boreal, temperate and tropical forests, respectively, and were 26-67% higher than those based on the original sampling depths used. We found significant positive correlations between ≤1 and ≤2 mm diameter roots and between ≤2 and ≤5 mm roots. The FRB estimates, standardized to the ≤2 mm diameter class, were 34-60% higher and 25-29% smaller than those standardized to the ≤1 mm and ≤5 mm diameter classes, respectively. The FRB of the understorey vegetation accounted for 31% of the total FRB in boreal forests and 20% in temperate forests. The results indicate that environmental factors (latitude, mean annual precipitation, elevation, temperature) or forest stand factors (life form, age, basal area, density) can not explain a significant amount of the variation in the total FRB and a maximum of 30% that in the FRB of trees at the stand level, whereas the mean basal area of the forest stand can explain 49% of the total FRB and 79% of the FRB of trees at the tree level.     

Uncertainty estimation of biomass expansion factors for Norway spruce in the Czech Republic

Nation wide estimates of the changes in forest biomass are needed for the greenhouse gas (GHG) reporting under the Climate Convention. The bases for national GHG reporting concerning forest sector are the national forest inventory (NFI) programmes. Since these programmes were mostly established for monitoring of timber resources, one of the current challenges for the NFIs is the development of methodology, such as biomass expansion factors (BEFs). The methodology for carbon stock change estimation should be transparent and verifiable, but this demand is not currently met due to the fact that the source data and uncertainty in the applied BEFs are not known. Here we developed BEFs with uncertainty estimation applicable to stand wise inventory of Norway spruce forests in the Czech Republic. BEFs were constructed, based on tree wise data from permanent research plots, by applying biomass and volume models to tree-level data. These BEFs were age-dependent and their uncertainty was sensitive to the dependencies among errors. Most of the uncertainty in the BEFs was due to uncertainty in the biomass and volume models applied.     

Interspecific competition impacts on the morphology and distribution of fine roots in European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.)

Morphology and vertical distribution patterns of spruce and beech live fine roots (diameter ≤2 mm) were studied using a soil core method in three comparable mature stands in the Solling: (1) pure beech, (2) pure spruce and (3) mixed spruce–beech. This study was aimed at determining the effects of interspecific competition on fine root structure and spatial fine root distribution of both species. A vertical stratification of beech and spruce fine root systems was found in the mixed stand due to a shift in beech fine roots from upper to lower soil layers. Moreover, compared to pure beech, a significantly higher specific root length (SRL, P<0.05) and specific surface area (SSA, P<0.05) were found for beech admixed with spruce (pure beech/mixed beech SRL 16.1–23.4 m g⁻¹, SSA 286–367 cm² g⁻¹). Both indicate a flexible ‘foraging’ strategy of beech tending to increase soil exploitation and space sequestration efficiency in soil layers less occupied by competitors. Spruce, in contrast, followed a more conservative strategy keeping the shallow vertical rooting and the root morphology quite constant in both pure and mixed stands (pure spruce/mixed spruce SRL 9.6/7.7 m g⁻¹, P>0.10; SSA 225/212 cm² g⁻¹, P>0.10). Symmetric competition belowground between mixed beech and spruce was observed since live fine roots of both species were under-represented compared to pure stand. However, the higher space sequestration efficiency suggests a higher competitive ability of beech belowground.