Temperature Sensitivity of Microbial Respiration of Fine Root Litter in a Temperate Broad-Leaved Forest

The microbial decomposition respiration of plant litter generates a major CO₂ efflux from terrestrial ecosystems that plays a critical role in the regulation of carbon cycling on regional and global scales. However, the respiration from root litter decomposition and its sensitivity to temperature changes are unclear in current models of carbon turnover in forest soils. Thus, we examined seasonal changes in the temperature sensitivity and decomposition rates of fine root litter of two diameter classes (0–0.5 and 0.5–2.0 mm) of Quercus serrata and Ilex pedunculosa in a deciduous broad-leaved forest. During the study period, fine root litter of both diameter classes and species decreased approximately exponentially over time. The Q₁₀ values of microbial respiration rates of root litter for the two classes were 1.59–3.31 and 1.28–6.27 for Q. serrata and 1.36–6.31 and 1.65–5.86 for I. pedunculosa. A significant difference in Q₁₀ was observed between the diameter classes, indicating that root diameter represents the initial substrate quality, which may determine the magnitude of Q₁₀ value of microbial respiration. Changes in these Q₁₀ values were related to seasonal soil temperature patterns; the values were higher in winter than in summer. Moreover, seasonal variations in Q₁₀ were larger during the 2-year decomposition period than the 1-year period. These results showed that the Q₁₀ values of fine root litter of 0–0.5 and 0.5–2.0 mm have been shown to increase with lower temperatures and with the higher recalcitrance pool of the decomposed substrate during 2 years of decomposition. Thus, the temperature sensitivity of microbial respiration in root litter showed distinct patterns according to the decay period and season because of the temperature acclimation and adaptation of the microbial decomposer communities in root litter.     

Disentangling the Litter Quality and Soil Microbial Contribution to Leaf and Fine Root Litter Decomposition Responses to Reduced Rainfall

Climate change-induced rainfall reductions in Mediterranean forests negatively affect the decomposition of plant litter through decreased soil moisture. However, the indirect effects of reduced precipitation on litter decomposition through changes in litter quality and soil microbial communities are poorly studied. This is especially the case for fine root litter, which contributes importantly to forests plant biomass. Here we analyzed the effects of long-term (11 years) rainfall exclusion (29% reduction) on leaf and fine root litter quality, soil microbial biomass, and microbial community-level physiological profiles in a Mediterranean holm oak forest. Additionally, we reciprocally transplanted soils and litter among the control and reduced rainfall treatments in the laboratory, and analyzed litter decomposition and its responses to a simulated extreme drought event. The decreased soil microbial biomass and altered physiological profiles with reduced rainfall promoted lower fine root—but not leaf—litter decomposition. Both leaf and root litter, from the reduced rainfall treatment, decomposed faster than those from the control treatment. The impact of the extreme drought event on fine root litter decomposition was higher in soils from the control treatment compared to soils subjected to long-term rainfall exclusion. Our results suggest contrasting mechanisms driving drought indirect effects on above-(for example, changes in litter quality) and belowground (for example, shifts in soil microbial community) litter decomposition, even within a single tree species. Quantifying the contribution of these mechanisms relative to the direct soil moisture-effect is critical for an accurate integration of litter decomposition into ecosystem carbon dynamics in Mediterranean forests under climate change.     

Nitrogen Controls on Fine Root Substrate Quality in Temperate Forest Ecosystems

Nitrogen controls on fine root substrate quality (that is, nitrogen and carbon-fraction concentrations) were assessed using nitrogen availability gradients in the Harvard Forest chronic nitrogen addition plots, University of Wisconsin Arboretum, Blackhawk Island, Wisconsin, and New England spruce-fir transect. The 27 study sites encompassed within these four areas collectively represented a wide range of nitrogen availability (both quantity and form), soil types, species composition, aboveground net primary production, and climatic regimes. Changes in fine root substrate quality among sites were most frequently and strongly correlated with nitrate availability. For the combined data set, fine root nitrogen concentration increased (adjusted R ² = 0.46, P < 0.0001) with increasing site nitrate availability. Fine root “extractive” carbon-fraction concentrations decreased (adjusted R ² = 0.32, P < 0.0002), “acid-soluble” compounds increased (adjusted R ² = 0.35, P < 0.0001), and the “acid-insoluble” carbon fraction remained relatively high and stable (combined mean of 48.7 ± 3.1% for all sites) with increasing nitrate availability. Consequently, the ratio of acid-insoluble C–total N decreased (adjusted R ² = 0.40, P < 0.0001) along gradients of increasing nitrate availability. The coefficients of determination for significant linear regressions between site nitrate availability and fine root nitrogen and carbon-fraction concentrations were generally higher for sites within each of the four study areas. Within individual study sites, tissue substrate quality varied between roots in different soil horizons and between roots of different size classes. However, the temporal variation of fine root substrate quality indices within specific horizons was relatively low. The results of this study indicate that fine root substrate quality increases with increasing nitrogen availability and thus supports the substrate quality component of a hypothesized conceptual model of nitrogen controls on fine root dynamics that maintains that fine root production, mortality, substrate quality, and decomposition increase with nitrogen availability in forest ecosystems in a manner that is analogous to foliage.     

Species and structural diversity of church forests in a fragmented Ethiopian Highland landscape

Question: Thousands of small isolated forest fragments remain around churches (“church forests”) in the almost completely deforested Ethiopian Highlands. We questioned how the forest structure and composition varied with altitude, forest area and human influence. Location: South Gondar, Amhara National Regional State, Northern Ethiopia. Methods: The structure and species composition was assessed for 810 plots in 28 church forests. All woody plants were inventoried, identified and measured (stem diameter) in seven to 56 10 m x 10‐m plots per forest. Results: In total, 168 woody species were recorded, of which 160 were indigeneous. The basal area decreased with tree harvest intensity; understorey and middle‐storey density (<5 cm DBH trees) decreased with grazing; overstorey density (>5 cm DBH trees) increased with altitude. The dominance of a small set of species increased with altitude and grazing intensity. Species richness decreased with altitude, mainly due to variation in the richness of the overstorey community. Moreover, species richness in the understorey decreased with grazing intensity. Conclusions: We show how tree harvesting intensity, grazing intensity and altitude contribute to observed variations in forest structure, composition and species richness. Species richness was, however, not related to forest area. Our study emphasizes the significant role played by the remaining church forests for conservation of woody plant species in North Ethiopian Highlands, and the need to protect these forests for plant species conservation purposes.     

Influence of Endomycorrhizal Infection on Root Morphology in a Micropropagated Woody Plant Species (Vitis vinifera L.)

The influence of vesicular-arbuscular (VA) endomycorrhizal infectionon root morphology and architecture of a woody micropropagatedplant, Vitis vinifera L., has been investigated using morphologicalanalysis, modelling and topological methods. Endomycorrhizaformation caused increases in lateral root number and consequentlytotal root length but did not alter the number of root axes.The rate of production of any order lateral roots was higherin mycorrhizal than non-mycorrhizal controls. The number offirst- and second-order laterals increased linearly with timein mycorrhizal plants whilst in control plants both fitted alogistic function. Topological analysis indicated similar patternsof root branching in the early stages of growth, but the rootsystem of non-mycorrhizal plants adopted a more herringbonepattern after 8 weeks, whereas that of mycorrhizal plants retaineda more dichotomous pattern with repeated bifurcation. Althoughthe root system pattern of non-mycorrhizal vines is more efficientin exploring soil, it is more expensive for the plant in termsof energy cost versus return benefit (nutrient acquisition).In contrast mycorrhizal plants develop a more economical rootsystem which is rendered more efficient by the direct role ofthe mycorrhizal fungus in assisting nutrient absorption. Vitis vinifera L., vine, root system, modelling, topology, vesicular-arbuscular mycorrhizae     

Root Morphology and Anchorage of Six Native Tree Species from a Tropical Montane Forest and an Elfin Forest in Ecuador

Root architecture of tree species was investigated at two different altitudes in tropical forests in Ecuador. Increasing altitude was accompanied by higher wind speeds and more shallow soils, while slope angles of both sites were comparable (20–50°). Three tree species typical for the montane forest at 1900 m (Graffenrieda emarginata (Ruiz & Pav.) Triana (Melastomataceae), Clethra revoluta (Ruiz & Pav.) Spreng. (Clethraceae), Vismia tomentosa Ruiz & Pav. (Clusiaceae)) and for the elfin forest at 3000 m (Weinmannia loxensis Harling (Cunoniaceae), Clusia spec. (Clusiacaea) Styrax foveolaria Perkins (Styraceae)) were examined. At 1900 m, 92% of the trees grew upright, in comparison to 52% at 3000 m. At 3000 m, 48% of the trees were inclined, lying or even partly uprooted. At this altitude, all trees with tap roots or with shoots connected by coarse rhizomes, 83% of the trees with stilt roots, and 50% of the trees in which stems or roots were supported by other trees grew upright, suggesting that these characteristics were relevant for tree stability. Root system morphology differed markedly between altitudes. In contrast to 1900 m, where 20% of structural roots originated in the deeper mineral soil, root origin at 3000 m was restricted to the forest floor. The mean ratio of root cross sectional area to tree height decreased significantly from 6.1 × 10⁻³ m² m⁻¹ at 1900 m to 3.2 × 10⁻³ m² m⁻¹ at 3000 m. The extent of root asymmetry increased significantly from 0.29 at 1900 m to 0.62 at 3000 m. This was accompanied by a significantly lower number of dominant roots at 3000 m (2.3 compared to 3.8 at 1900 m). In conclusion, native tree species growing in tropical montane and elfin forests show a variety of root traits that improve tree stability. Root system asymmetry is less important for tree stability where anchorage is provided by a deep and solid root–soil plate. When deep rooting is impeded, root traits improving the horizontal extension of the root–soil plate are more pronounced or occur more frequently. Furthermore, mutual mechanical support of roots and stems of neighboring trees seems to be an appropriate mechanism to provide anchorage in soils with low bulk density and in environments with high wind speeds.     

Seed Ecology of Woody Species in a Caribbean Dry Forest

Peak flowering activity among woody species in the tropical dry forests of St. John, U.S. Virgin Islands, coincided with the brief spring rainy season but continued at moderate levels for six months, abating with the autumn rains. Fruit maturation showed a major peak in the long winter dry season and a minor crest during the summer dry season. Seeds of wind-dispersed species disseminated mainly during the winter dry season, while animal dispersal of seeds (74% of all woody species) followed the bimodal pattern (for wet and dry seasons) described for the community as a whole. Under shadehouse conditions, most dry forest tree species germinated well (> 80%) and emerged promptly (within four weeks of planting) and synchronously (90% emergence within a four-week interval). Nine of 29 species tested in the shadehouse manifested dormancy of at least six weeks. Seed germinability varied among tree species, and the viability of most species began to decline following six months of dry storage. Few species retained high germinability after nine months of dry storage. The species composition of soil seed banks did not correspond closely with above-ground communities on three forested sites of varying stand age. In the youngest stand (35 years old), dominated by the weedy, arborescent legume Leucaena leucocephala, the soil seed bank was also dominated by this species, but no seeds of any other tree species were found in the soil samples. Seeds of native trees were scarcely encountered (only one indigenous species) in soil seed bank samples of three forest sites. Local seed rain from less disturbed forest may not be sufficient for prompt recovery of the dry forest community on degraded sites.     

Litter Decomposition in Four Woody Species in a Mediterranean Climate: Weight Loss, N and P Dynamics

Decomposition dynamics in leaves and needles of two Mediterraneanshrubs and two pine species growing in the Sierra de Filabres(Almería, Spain) was investigated during 2 years usingthe litter bag technique. The species studied are representativeof the vegetation of the study area and differ greatly in theirfoliar traits. Results are discussed in relation to the initiallitter quality (C, N and P) and through the application of theexponential decay model. The mass lost at the end of study variedin the order: Pinus pinaster < Pinus nigra < Cistus laurifolius< Adenocarpus decorticans. Differences in annual rates ofdecomposition among species are consistent with the particularchemical and structural attributes of their leaves. The massof decomposing litter remaining after 2 years was positivelyassociated with the initial C:N ratio. Weight loss and nutrientrelease were fastest in the leguminous A. decorticans. The resultssuggest the importance of both structure and elemental concentrationof initial litter for decomposition dynamics in Mediterraneanspecies. Copyright 2000 Annals of Botany Company Adenocarpus decorticans Boiss., C:N ratio, Cistus laurifolius L., litter decomposition, litter quality, Mediterranean environments, nutrient dynamics, Pinus pinaster Aiton, Pinus nigra Arnold, single exponential model, Southern Spain     

Spatial Variability in Litterfall,Litter Standing Crop and Litter Quality in a Tropical Rain Forest Region

Understanding the spatial variability in plant litter processes is essential for accurate comprehension of biogeochemical cycles and ecosystem function. We assessed spatial patterns in litter processes from local to regional scales, at sites throughout the wet tropical rain forests of northern Australia. We aimed to determine the controls (e.g., climate, soil, plant community composition) on annual litter standing crop, annual litterfall rate and in situ leaf litter decomposability. The level of spatial variance in these components, and leaf litter N, P, Ca, lignin, α‐cellulose and total phenolics, was determined from within the scale of subregion, to site (1 km transects) to local/plot (~30 m²). Overall, standing crop was modeled with litterfall and its chemical composition, in situ decomposability, soil Na, and topography (r² = 0.69, 36 plots). Litterfall was most closely aligned with plant species richness and stem density (negative correlation); leaf decomposability with leaf‐P and lignin, soil Na, and dry season moisture (r² = 0.89, 40 plots). The predominant scale of variability in litterfall rates was local (plot), while litter standing crop and α‐cellulose variability was more evenly distributed across spatial scales. Litter decomposability, N, P and phenolics were more aligned with subregional differences. Leaf litter C, lignin and Ca varied most at the site level, suggesting more local controls. We show that variability in litter quality and decomposability are more easily accounted for spatially than litterfall rates, which vary widely over short distances possibly in response to idiosyncratic patterns of disturbance.     

南亚热带森林木本植物种子萌发和储存

研究了南亚热带森林 90种木本植物的种子和果实的大小、含水量等特征及种子萌发 ,并对其中有足够萌发个体的种 (45种 )进行了萌发分析和对有足够萌发个体和足够种子的种 (41种 )进行了种子或果实储存对萌发影响的研究。肉质果实种子和肉质种子萌发率较干燥种子高 ,两者差异显著。大多数种播种 6 0d内萌发。肉质果实的种子和肉质种子较干燥种子的休眠期长 ,留土萌发种子较出土萌发种子休眠期长。大种子种与小种子种之间、夏季成熟的肉质果实的种子与冬季和春季成熟的肉质果实的种子之间休眠期长短差异不显著。萌发率与新鲜种子含水量无显著的相关 ,但种子中水分含量的减少明显影响萌发率 ,尤其是在种子自然干燥的最初 3d。当种子的水分含量减至 2 0 % (约在种子自然干燥 10～ 14d)时 ,种子萌发率降至很低。萌发率随储存时间 (密封 ,(4± 1)℃ )而下降 ,但储存一个月下降不大 (- 8.3% ) ,储存 3个月后 ,显著下降。果实储存与种子储存之间、大种子种与小种子种储存之间、肉质果实的种子和肉质种子储存与干燥种子储存之间 ,萌发率差异一般不显著。     

南亚热带森林木本植物种子萌发和储存

研究了南亚热带森林90种木本植物的种子和果实的大小、含水量等特征及种子萌发,并对其中有足够萌发个体的种(45种)进行了萌发分析和对有足够萌发个体和足够种子的种(41种)进行了种子或果实储存对萌发影响的研究.肉质果实种子和肉质种子萌发率较干燥种子高,两者差异显著.大多数种播种60 d内萌发.肉质果实的种子和肉质种子较干燥种子的休眠期长,留土萌发种子较出土萌发种子休眠期长.大种子种与小种子种之间、夏季成熟的肉质果实的种子与冬季和春季成熟的肉质果实的种子之间休眠期长短差异不显著.萌发率与新鲜种子含水量无显著的相关,但种子中水分含量的减少明显影响萌发率,尤其是在种子自然干燥的最初3 d.当种子的水分含量减至20%(约在种子自然干燥10～14 d)时,种子萌发率降至很低.萌发率随储存时间(密封,(4±1) ℃)而下降,但储存一个月下降不大(-8.3%),储存3个月后,显著下降.果实储存与种子储存之间、大种子种与小种子种储存之间、肉质果实的种子和肉质种子储存与干燥种子储存之间,萌发率差异一般不显著.     

Coarse Woody Debris Stimulates Soil Enzymatic Activity and Litter Decomposition in an Old-Growth Temperate Forest of Patagonia, Argentina

In most temperate forest ecosystems, tree mortality over time generates downed logs that accumulate as coarse woody debris (CWD) on the forest floor. These downed logs and trunks have important recognized ecosystem functions including habitat for different organisms and long-term organic C storage. Due to its recalcitrant chemical composition and slow decomposition, CWD can also have direct effects on ecosystem carbon and nutrient turnover. CWD could also cause changes indirectly through the physical and chemical alterations that it generates, although it is not well-understood how important these indirect effects could be for ecosystem processes and soil biogeochemistry. We hypothesized that in an old-growth mature forest, CWD affects carbon and nutrient cycles through its “proximity effects”, meaning that the forest floor near CWD would have altered soil biotic activity due to the environmental and biogeochemical effects of the presence of CWD. We conducted our study in an old-growth southern beech temperate forest in Patagonia, Argentina, where we estimated and classified the distribution and mass, nutrient pools and decay stage of CWD on the forest floor, and evaluated its impact on litter decomposition, soil mites and soil enzymatic activity of carbon and phosphorus-degrading enzymes. We demonstrate here that CWD in this ecosystem represents an important organic carbon reservoir (85 Mg ha^?1) and nitrogen pool (0.42 Mg ha^?1), similar in magnitude to other old-growth forests of the Northern Hemisphere. In addition, we found significant proximity effects of CWD, with increased C-degrading soil enzyme activity, decreased mite abundance, and more rapid litter decomposition beneath highly decayed CWD. Considered at the ecosystem scale in this forest, the removal of CWD could cause a decrease of 6% in soil enzyme activity, particularly in the summer dry season, and nearly 15% in annual litter decomposition. We conclude that beyond the established importance of CWD as a long-term carbon reservoir and habitat, CWD contributes functionally to the forest floor by influencing the spatial heterogeneity of microbial activity and carbon and nutrient turnover. These proximity effects demonstrate the importance of maintenance of this ecosystem component and should be taken into consideration for management decisions pertaining to carbon sequestration and functional diversity in natural forest ecosystems.     

桢楠种源幼苗细根形态和生物量研究

为了解桢楠(Phoebe zhennan)不同种源细根形态和生物量分配的差异,采用全根调查的方法,对桢楠自然分布区13个种源2.5年生幼苗的细根形态和生物量进行了研究。结果表明,桢楠种源间各级细根的平均直径、总根长和表面积差异显著,在种源内细根的平均直径随根序的增加而增加,但根序间总根长和表面积差异规律不明显。根序生物量分配随根序增加而增加,1~4级根生物量分配分别为6.33%、14.47%、25.03%和54.17%。通过综合评价,以HT、LF、ES和WC种源的根系最优,具有较高的生长潜力。     

森林生态系统细根周转规律及影响因素

根系周转是陆地生态系统碳循环的关键过程, 对研究土壤碳库变化及全球气候变化均具有重要意义。然而由于根系周转率的测量计算方法较多, 不同方法得出的结果差异较大, 且目前对全球区域尺度上森林生态系统根系周转的研究还不够充分, 使得全球森林生态系统根系周转变化规律仍不清楚。该研究通过收集文献数据并统一周转率计算方法, 对全球5种森林类型的细根周转空间格局进行整合, 同时结合土壤理化性质和气候数据, 得出影响森林生态系统细根周转的因子。结果表明, 不同森林类型细根周转率存在显著差异, 且随着纬度的升高逐渐降低; 森林生态系统细根周转率与年平均温度和年平均降水量呈正相关; 森林生态系统细根周转率与土壤有机碳含量呈正相关但与土壤pH值呈负相关。该研究为揭示森林生态系统细根周转规律及机制提供了科学依据。     

Small Mammal Responses to Fine Woody Debris and Forest Fuel Reduction in Southwest Oregon

Abstract: Despite its importance for wildlife, most forests in the Pacific Northwest contain low volumes of large downed wood compared to fine woody debris (FWD). We used a replicated experiment to compare short-term responses of deer mice (Peromyscus maniculatus) and western red-backed voles (Clethrionomys californicus) among 3 arrangements of FWD: piled, lopped and scattered, and pile burning, a commonly used method of fuel reduction in commercial Douglas fir (Pseudotsuga menziesii) forests in southwest Oregon, USA. We assessed habitat use, density, and survival of mice and voles during 2 consecutive summers (Jun—Aug 1999 and 2000). Both mice and voles used FWD cover disproportionately from its availability, and they differed in their responses to specific FWD arrangements. Mice used piled FWD (proportional use = 37.0%, 90% CI = 33.0–44.0) 43% more than expected (26.0). Number of mice captured (x̄ = 1.9 mice, 90% CI = 1.5–2.5) and index of home range size (x̄ = 4.8 m, 90% CI = 0.7–8.9) at individual FWD piles decreased up to 16% and increased up to 50%, respectively, for each 1-m increase in distance from piles. Voles used all FWD cover classes in proportion to availability, but number of voles captured increased slightly (x̄ = 0.016 voles/m, 90% CI = 0.001–0.031) for each 1-m increase in distance from piles. Piled FWD had no discernable effect on population density and apparent survival of mice, but analyses had low power (0.25, 0.67). Our results suggest that piling FWD would benefit deer mice, whereas lopped and scattered FWD might benefit voles. Thus, a combination of methods to reduce fire risk should be considered to accommodate multiple small mammal species. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):625–632; 2008)     

凋落物对土壤酸化的缓冲及其对根系生长的影响

凋落物层是森林生态系统中一个十分活跃的界面,一方面环境因素影响凋落物的积累和分解,另一方面凋落物的动态反过来又影响到系统内的水热收支平衡、土壤的理化特性以至森林生产力。值得注意的是,在高度工业化的现代社会,大气污染作为一种新生的环境因子不同程度地作用于森林生态系统,凋落物层则是大气污染物(如酸雨)作用于森林土壤亚系     

Woody Species Diversity in Forest Plantations in a Mountainous Region of Beijing,China: Effects of Sampling Scale and Species Selection

The role of forest plantations in biodiversity conservation has gained more attention in recent years. However, most work on evaluating the diversity of forest plantations focuses only on one spatial scale; thus, we examined the effects of sampling scale on diversity in forest plantations. We designed a hierarchical sampling strategy to collect data on woody species diversity in planted pine (Pinus tabuliformis Carr.), planted larch (Larix principis-rupprechtii Mayr.), and natural secondary deciduous broadleaf forests in a mountainous region of Beijing, China. Additive diversity partition analysis showed that, compared to natural forests, the planted pine forests had a different woody species diversity partitioning pattern at multi-scales (except the Simpson diversity in the regeneration layer), while the larch plantations did not show multi-scale diversity partitioning patterns that were obviously different from those in the natural secondary broadleaf forest. Compare to the natural secondary broadleaf forests, the effects of planted pine forests on woody species diversity are dependent on the sampling scale and layers selected for analysis. Diversity in the planted larch forest, however, was not significantly different from that in the natural forest for all diversity components at all sampling levels. Our work demonstrated that the species selected for afforestation and the sampling scales selected for data analysis alter the conclusions on the levels of diversity supported by plantations. We suggest that a wide range of scales should be considered in the evaluation of the role of forest plantations on biodiversity conservation.     

Fine Root Distribution in a Lower Montane Rain Forest of Panama

In a Panamanian lower montane rain forest we: (1) analyzed the vertical and horizontal distribution of fine roots; and (2) assessed the relationship of fine root mass to thickness of the soil organic layer, soil pH, and soil-extractable nitrogen. The soil in the study area has developed on volcanic ash deposits and was classified as Hapludand. In randomly distributed samples, the median fine root mass (biomass and necromass, diam ≤ 2 mm) to a depth of 100 cm mineral soil was 544 g/m², 41 percent of which was found in the organic layer. Fine root mass was approximately twice as high in the vicinity of stems of the tree species Oreomunnea mexicana (1069 g/m²) and the palm species Colpothrinax aphanopetala (1169 g/m²) and was associated with thick organic layers. The median thickness of the soil organic layer in a larger random sample ( N = 64) was 8 cm with a considerable variation (interquartile range: 7 cm). In these samples, the density of fine root biomass was correlated with the concentration of extractable nitrogen ( r = 0.33, P = 0.011), and on an areal basis, fine root biomass in the organic layer increased with increasing thickness of the organic layer ( r = 0.63, P < 0.001) and decreasing pH_KCl ( r =−0.33, P < 0.01). Fine root biomass in the upper mineral soil did not show significant correlations with any of the studied parameters.