Litter decomposition contrasts in second- and old-growth Douglas-fir forests of the Pacific Northwest, USA

Litterfall and its subsequent decomposition are important feedback mechanisms in the intrasystem cycling of nutrients in forest ecosystems. The amount of litterfall and the rate of decomposition are expected to vary with stand age and climate. Over a 2-year period, decomposition of five litter types were measured in two second-growth forest stands and one old-growth stand in the Cascade Mountains of southern Washington state, USA. Both second-growth stands were dominated by Douglas-fir [Pseudotsuga menziesii (Mirb.,) Franco] but one had a significant proportion of red alder (Alnus rubra Bong.), a nitrogen (N) fixer. The old-growth stand was dominated by Douglas-fir and western hemlock [Tsuga heterophylla (Raf.) Sarg.]. All stands had a relatively shallow layer of forest floor mass. The five litter types were placed in each stand to evaluate decomposition patterns. Despite significant differences in stand age, microclimate and mean residence times for carbon (C) and N, the rates of litter mass loss varied only slightly between sites. The relative order of species litter mass loss was: vine maple ≫ salal = western hemlock > Douglas-fir (from the youngest stand) > Douglas-fir (from the N rich stand with red alder). The initial litter lignin concentration, not lignin:N, was the primary determinant of decomposition rates, although the initial N concentration was the predictor for mass loss after 2 years in the N rich Douglas-fir-alder stand. All litter types showed immobilization of N for nearly 2 years. Data for Douglas-fir litter suggest that higher levels of N may retard decomposition of tissues with greater amounts of lignified material. The retention of N by the litter appeared influenced by the nutrient capital of the stands as well as the forest floor C:N ratio. Decomposition was minimal during the cold winter months, but displayed a definitive peak period during early Fall with wet weather, warm soils, and fungal activity. Thus, long-term climatic change effects on forest floor C storage may depend more on changes in seasonality of precipitation changes than just temperature changes.     

Biomass, morphology and nutrient contents of fine roots in four Norway spruce stands

Fine root systems may respond to soil chemical conditions, but contrasting results have been obtained from field studies in non-manipulated forests with distinct soil chemical properties. We investigated biomass, necromass, live/dead ratios, morphology and nutrient concentrations of fine roots (<2 mm) in four mature Norway spruce (Picea abies [L.] Karst.) stands of south-east Germany, encompassing variations in soil chemical properties and climate. All stands were established on acidic soils (pH (CaCl₂) range 2.8–3.8 in the humus layer), two of the four stands had molar ratios in soil solution below 1 and one of the four stands had received a liming treatment 22 years before the study. Soil cores down to 40 cm mineral soil depth were taken in autumn and separated into four fractions: humus layer, 0–10 cm, 10–20 cm and 20–40 cm. We found no indications of negative effects of N availability on fine root properties despite large variations in inorganic N seepage fluxes (4–34 kg N ha⁻¹ yr⁻¹), suggesting that the variation in N deposition between 17 and 26 kg N ha⁻¹ yr⁻¹ does not affect the fine root system of Norway spruce. Fine root biomass was largest in the humus layer and increased with the amount of organic matter stored in the humus layer, indicating that the vertical pattern of fine roots is largely affected by the thickness of this horizon. Only two stands showed significant differences in fine root biomass of the mineral soil which can be explained by differences in soil chemical conditions. The stand with the lowest total biomass had the lowest Ca/Al ratio of 0.1 in seepage, however, Al, Ca, Mg and K concentrations of fine roots were not different among the stands. The Ca/Al ratio in seepage might be a less reliable stress parameter because another stand also had Ca/Al ratios in seepage far below the critical value of 1.0 without any signs of fine root damages. Large differences in the live/dead ratio were positively correlated with the Mn concentration of live fine roots from the mineral soil. This relationship was attributed to faster decay of dead fine roots because Mn is known as an essential element of lignin degrading enzymes. It is questionable if the live/dead ratio can be used as a vitality parameter of fine roots since both longevity of fine roots and decay of root litter may affect this parameter. Morphological properties were different in the humus layer of one stand that was limed in 1983, indicating that a single lime dose of 3–4 Mg ha⁻¹ has a long-lasting effect on fine root architecture of Norway spruce. Almost no differences were found in morphological properties in the mineral soil among the stands, but vertical patterns were apparently different. Two stands with high base saturation in the subsoil showed a vertical decrease in specific root length and specific root tip density whereas the other two stands showed an opposite pattern or no effect. Our results suggest that proliferation of fine roots increased with decreasing base saturation in the subsoil of Norway spruce stands.     

Fine Root Production across a Primary Successional Ecosystem Chronosequence at Mt. Shasta,California

Estimating changes in belowground biomass and production is essential for understanding fundamental patterns and processes during ecosystem development. We examined patterns of fine root production, aboveground litterfall, and forest floor accumulation during forest primary succession at the Mt. Shasta Mudflows ecosystem chronosequence. Fine root production was measured using the root ingrowth cores method over 1 year, and aboveground litterfall was collected over 2 years. Fine root production increased significantly with ecosystem age, but only the youngest ecosystem was significantly different from all of the older ecosystems. Root production was 44.5 ± 13.3, 168.3 ± 20.6, 190.5 ± 33.8, and 236.3 ± 65.4 g m⁻² y⁻¹ in the 77, 255, 616, and >850-year-old ecosystems, respectively. Generally, aboveground litterfall and forest floor accumulation did not follow the same pattern as root production. The relative contribution of fine root production to total fine detrital production increased significantly with ecosystem age, from 14 to 49%, but only the youngest ecosystem was significantly different from all of the older ecosystems. Fine root production was significantly correlated with some measures of soil fertility but was not correlated with leaf or total litterfall, or forest floor accumulation. It was best predicted by soil N concentration alone, but this relationship may not be causal, as soil N concentration was also correlated with ecosystem age. For the oldest ecosystem, fine root production was also measured using the sequential intact cores/compartment-flow model method, and the difference between the two estimates was not significant. Our study suggests that the relative contribution of fine roots to fine detrital production, and hence to soil organic matter accumulation, may increase during forest primary succession.     

Fine Root Dynamics and Forest Production Across a Calcium Gradient in Northern Hardwood and Conifer Ecosystems

Losses of soil base cations due to acid rain have been implicated in declines of red spruce and sugar maple in the northeastern USA. We studied fine root and aboveground biomass and production in five northern hardwood and three conifer stands differing in soil Ca status at Sleepers River, VT; Hubbard Brook, NH; and Cone Pond, NH. Neither aboveground biomass and production nor belowground biomass were related to soil Ca or Ca:Al ratios across this gradient. Hardwood stands had 37% higher aboveground biomass (P = 0.03) and 44% higher leaf litter production (P < 0.01) than the conifer stands, on average. Fine root biomass (<2 mm in diameter) in the upper 35 cm of the soil, including the forest floor, was very similar in hardwoods and conifers (5.92 and 5.93 Mg ha⁻¹). The turnover coefficient (TC) of fine roots smaller than 1 mm ranged from 0.62 to 1.86 y⁻¹ and increased significantly with soil exchangeable Ca (P = 0.03). As a result, calculated fine root production was clearly higher in sites with higher soil Ca (P = 0.02). Fine root production (biomass times turnover) ranged from 1.2 to 3.7 Mg ha⁻¹ y⁻¹ for hardwood stands and from 0.9 to 2.3 Mg ha⁻¹ y⁻¹ for conifer stands. The relationship we observed between soil Ca availability and root production suggests that cation depletion might lead to reduced carbon allocation to roots in these ecosystems.     

Fine Root Biomass,Production, Turnover Rates,and Nutrient Contents in Boreal Forest Ecosystems in Relation to Species,Climate, Fertility,and Stand Age: Literature Review and Meta-Analyses

Fine roots <2 mm in diameter play a key role in regulating the biogeochemical cycles of ecosystems and are important to our understanding of ecosystem responses to global climate changes. Given the sensitivity of fine roots, especially in boreal region, to climate changes, it is important to assess whether and to what extent fine roots in this region change with climates. Here, in this synthesis, a data set of 218 root studies were complied to examine fine root patterns in the boreal forest in relation to site and climatic factors. The mean fine root biomass in the boreal forest was 5.28 Mg ha^?1, and the production of fine roots was 2.82 Mg ha^?1 yr^?1, accounting for 32% of annual net primary production of the boreal forest. Fine roots in the boreal forest on average turned over 1.07 times per year. Fine roots contained 50.9 kg ha^?1 of nitrogen (N) and 3.63 kg ha^?1 of phosphorous (P). In total, fine roots in the boreal forest ecosystems contain 6.1 × 10⁷ Mg N and 4.4×10⁶Mg P pools, respectively, about 10% of the global nutrients of fine roots. Fine root biomass, production, and turnover rate generally increased with increasing mean annual temperature and precipitation. Fine root biomass in the boreal forest decreased significantly with soil N and P availability. With increasing stand age, fine root biomass increased until about 100 years old for forest stands and then leveled off or decreased thereafter. These results of meta analysis suggest that environmental factors strongly influence fine root biomass, production, and turnover in boreal forest, and future studies should place a particular emphasis on the root-environment relationships.     

Nutrient cycling in Pinus sylvestris stands in eastern Finland

Nutrient cycling within three Pinus sylvestris stands was studied in eastern Finland. The aim of the study was to determine annual fluxes and distribution of N, P, K, Ca, Mg, Zn, Fe, B, and Al in the research stands. Special emphasis was put on determining the importance of different fluxes, especially the internal cycle within the trees in satisfying the tree nutrient requirements for biomass production. The following nutrient fluxes were included, input; free precipitation and throughfall, output; percolation through soil profile, biological cycle; nutrient uptake from soil, retranslocation within trees, return to soil in litterfall, release by litter decomposition. The distribution of nutrients was determined in above- and belowground tree compartments, in ground and field vegetation, and in soil.The nitrogen use efficiencies were 181, 211 and 191 g of tree aboveground dry matter produced per g of N supplied by uptake and retranslocation in the sapling, pole stage and mature stands, respectively. Field vegetation was more efficient in nitrogen use than trees. Stand belowground/aboveground and fine root/coarse root biomass ratios decreased with tree age. With only slightly higher fine root biomass, almost three times more nitrogen had to be taken-up from soil for biomass production in the mature stand than in the sapling stand.The annual input-output balances of most nutrients were positive; throughfall contained more nutrients than was lost in mineral soil leachate. The sulphate flux contributed to the leaching of cations, especially magnesium, from soil in the mature stand.Retranslocation supplied 17–42% of the annual N, P and K requirements for tree aboveground biomass production. Precipitation and throughfall were important in transferring K and Mg, and also N in the sapling stand. Litterfall was an important pathway for N, Ca, Mg and micro nutrients, especially in the oldest stands.     

Biomass and morphology of fine roots in temperate broad-leaved forests differing in tree species diversity: is there evidence of below-ground overyielding?

Biodiversity effects on ecosystem functioning in forests have only recently attracted increasing attention. The vast majority of studies in forests have focused on above-ground responses to differences in tree species diversity, while systematic analyses of the effects of biodiversity on root systems are virtually non-existent. By investigating the fine root systems in 12 temperate deciduous forest stands in Central Europe, we tested the hypotheses that (1) stand fine root biomass increases with tree diversity, and (2) ‘below-ground overyielding’ of species-rich stands in terms of fine root biomass is the consequence of spatial niche segregation of the roots of different species. The selected stands represent a gradient in tree species diversity on similar bedrock from almost pure beech forests to medium-diverse forests built by beech, ash, and lime, and highly-diverse stands dominated by beech, ash, lime, maple, and hornbeam. We investigated fine root biomass and necromass at 24 profiles per stand and analyzed species differences in fine root morphology by microscopic analysis. Fine root biomass ranged from 440 to 480 g m⁻² in the species-poor to species-rich stands, with 63–77% being concentrated in the upper 20 cm of the soil. In contradiction to our two hypotheses, the differences in tree species diversity affected neither stand fine root biomass nor vertical root distribution patterns. Fine root morphology showed marked distinctions between species, but these root morphological differences did not lead to significant differences in fine root surface area or root tip number on a stand area basis. Moreover, differences in species composition of the stands did not alter fine root morphology of the species. We conclude that ‘below-ground overyielding’ in terms of fine root biomass does not occur in the species-rich stands, which is most likely caused by the absence of significant spatial segregation of the root systems of these late-successional species.     

Root Growth and Recovery in Temperate Broad-Leaved Forest Stands Differing in Tree Species Diversity

In contrast to studies on aboveground processes, the effect of species diversity on belowground productivity and fine-root regrowth after disturbance is still poorly studied in forests. In 12 old-growth broad-leaved forest stands, we tested the hypotheses that (i) the productivity and recovery rate (regrowth per standing biomass) of the fine-root system (root diameter < 2 mm) increase with increasing tree species diversity, and that (ii) the seasonality of fine-root biomass and necromass is more pronounced in pure than in tree species-rich stands as a consequence of non-synchronous root biomass peaks of the different species. We investigated stands with 1, 3, and 5 dominant tree species growing under similar soil and climate conditions for changes in fine-root biomass and necromass during a 12-month period and estimated fine-root productivity with two independent approaches (ingrowth cores, sequential coring). According to the analysis of 360 ingrowth cores, fine-root growth into the root-free soil increased with tree species diversity from 72 g m⁻² y⁻¹ in the monospecific plots to 166 g m⁻² y⁻¹ in the 5-species plots, indicating an enhanced recovery rate of the root system after soil disturbance with increasing species diversity (0.26, 0.34, and 0.51 y⁻¹ in 1-, 3-, and 5-species plots, respectively). Fine-root productivity as approximated by the sequential coring data also indicated a roughly threefold increase from the monospecific to the 5-species stand. We found no indication of a more pronounced seasonality of fine-root mass in species-poor as compared to species-rich stands. We conclude that species identification on the fine root level, as conducted here, may open new perspectives on tree species effects on root system dynamics. Our study produced first evidence in support of the hypothesis that the fine-root systems of more diverse forest stands are more productive and recover more rapidly after soil disturbance than that of species-poor forests.     

Fine root vertical distribution and temporal dynamics in mature stands of two enset (Enset ventricosum Welw Cheesman) clones

Quantification of the role of fine roots in the biological cycle of nutrients necessitates understanding root distribution, estimating root biomass, turnover rate and nutrient concentrations, and the dynamics of these parameters in perennial systems. Temporal dynamics, vertical distribution, annual production and turnover, and nitrogen use of fine roots (≤2 mm in diameter) were studied in mature (5-year-old) stands of two enset (Ensete ventricosum) clones using the in-growth bag technique. Live fine root mass generally decreased with increasing depth across all seasons except the dry period. Except for the dry period, more than 70% of the fine root mass was in the above 0-20 cm depth, and the fine root mass in the upper 0–10 cm depth was significantly higher than in the lowest depth (20–30 cm). Live fine root mass showed a seasonal peak at the end of the major rainy season but fell to its lowest value during the dry or short rainy season. The difference between the peak and low periods were significant (p ≤ 0.05). Fine root nitrogen (N) use showed significant seasonal variation where the mean monthly fine root N use was highest during the major rainy season. There were significant effects on N use due to depths and in-growth periods, but not due to clones. Enset fine root production and turnover ranged from 2,339 to 2,451 kg ha⁻¹ year⁻¹ and from 1.55 to 1.80 year⁻¹, respectively. Root N return, calculated from fine root turnover, was estimated at 64–65 kg ha⁻¹ year⁻¹. Fine root production, vertical distribution and temporal dynamics may be related to moisture variations and nutrient (N) fluxes among seasons and along the soil depth. The study showed that fine root production and turnover can contribute considerably to the carbon and nitrogen economy of mature enset plots.     

杉木林年龄序列地下碳分配变化

  森林地下碳分配在森林碳平衡和碳吸存中具有重要作用, 而揭示人工林生长过程中地下碳分配变化对于人工林碳汇估算和碳汇管理等有重要意义。通过采用年龄序列方法研究了杉木(Cunninghamia lanceolata)林生长过程中地下碳分配变化特点。年龄序列为福建省南平7 a生(幼龄林)、16 a生(中龄林)、21 a生(近熟林)、41 a生(成熟林)和88 a生(老龄林)的杉木林。细根净生产力测定采用连续土芯法, 根系呼吸测定采用壕沟法, 生物量增量测定采用异速生长方程, 地上年凋落物量采用凋落物收集框测定。结果表明: 杉木林细根净生产力在中龄林前没有显著差异, 维持在较高水平; 但此后则显著下降。细根净生产力/地上凋落物量比值随林龄增加而显著下降。老龄林的根系呼吸显著低于其它林龄林分, 根系呼吸与细根生物量间呈显著线性相关。中龄林和近成熟林的地下碳分配(Total belouground carbon allocation, TBCA)显著高于幼龄林和成熟林, 而老龄林的则最低。中龄林、近成熟林和成熟林的地上部分净生产力/TBCA比值显著高于幼龄林和老龄林, 而杉木林的根系碳利用效率(RCUE)则呈现出随林龄增加而降低的趋势。     

Biomass and net production in a bambooPhyllostachys bambusoides stand

Biomass and net production were measured in aPhyllostachys bambusoides stand in Kyoto Prefecture, central Japan, which had carried out gregarious flowering in 1969 and has been recovering vegetatively. The culm density fluctuated around an average value of 12 040 ha⁻¹ during the research period (1985–91). Annual recruirment and mortality rates of culms were 1340 and 1133 ha⁻¹, respectively. The mean diameter at breast height increased from 7.28 cm in 1985 to 8.68 cm in 1991, and the biomass of culms increased from 71.3 to 111.6t ha⁻¹ over the same time period. Branch and leaf biomasses were almost constant, 10.0 and 9.4t ha⁻¹ on average, respectively. The leaf area index of the stand was 11.6 ha ha⁻¹, which is one of the largest values found in Japanese forests. The belowground biomass of 32.6t ha⁻¹ for rhizomes and 14.8t ha⁻¹ for fine roots resulted in the smaller ratio of aboveground parts to the root system (2.38) than those determined for forest stands. The amount of litterfall, excluding culms and large branches, was large (9.13t ha⁻¹ year⁻¹), corresponding to those measured in equatorial stands. The aboveground net production was 24.6t ha⁻¹ year⁻¹, larger than the average value reported for forest stands under similar weather conditions.     

Stand fine root biomass and fine root morphology in old-growth beech forests as a function of precipitation and soil fertility

Only very limited information exists on the plasticity in size and structure of fine root systems, and fine root morphology of mature trees as a function of environmental variation. Six northwest German old-growth beech forests (Fagus sylvatica L.) differing in precipitation (520 – 1030 mm year^–1) and soil acidity/fertility (acidic infertile to basic fertile) were studied by soil coring for stand totals of fine root biomass (0–40 cm plus organic horizons), vertical and horizontal root distribution patterns, the fine root necromass/biomass ratio, and fine root morphology (root specific surface area, root tip frequency, and degree of mycorrhizal infection). Stand total of fine root biomass, and vertical and horizontal fine root distribution patterns were similar in beech stands on acidic infertile and basic fertile soils. In five of six stands, stand fine root biomass ranged between 320 and 470 g m^–2; fine root density showed an exponential decrease with soil depth in all profiles irrespective of soil type. An exceptionally small stand fine root biomass (<150 g m^–2) was found in the driest stand with 520 mm year^–1 of rainfall. In all stands, fine root morphological parameters changed markedly from the topsoil to the lower profile; differences in fine root morphology among the six stands, however, were remarkably small. Two parameters, the necromass/biomass ratio and fine root tip density (tips per soil volume), however, were both much higher in acidic than basic soils. We conclude that variation in soil acidity and fertility only weakly influences fine root system size and morphology of F. sylvatica, but affects root system structure and, probably, fine root mortality. It is hypothesized that high root tip densities in acidic infertile soils compensate for low nutrient supply rates, and large necromasses are a consequence of adverse soil chemical conditions. Data from a literature survey support the view that rainfall is another major environmental factor that influences the stand fine root biomass of F. sylvatica.     

Shifts in litterfall and dominant nitrogen sources after expansion of shrub thickets

Woody encroachment into herbaceous ecosystems is emerging as an important ecological response to global change. A primary concern is alterations in C and N cycling and associated variations across a variety of ecosystems. We quantified seasonal variation in litterfall and litter N concentration in Morella cerifera shrub thickets to assess changes in litterfall and associated N input after shrub expansion on an Atlantic coast barrier island. We also used the natural abundance of ¹⁵N to estimate the proportion of litterfall N originating from symbiotic N fixation. Litterfall for shrub thickets ranged from 8,991 ± 247 to 3,810 ± 399 kg ha⁻¹ year⁻¹ and generally declined with increasing thicket age. Litterfall in three of the four thickets exceeded previous estimates of aboveground annual net primary production in adjacent grasslands by 300–400%. Leaf N concentration was also higher after shrub expansion and, coupled with low N resorption efficiency and high litterfall, resulted in a return of as much as 169 kg N ha⁻¹ year⁻¹ to the soil. We estimated that ∼70% of N returned to the soil was from symbiotic N fixation resulting in an ecosystem input of between 37 and 118 kg ha⁻¹ year⁻¹ of atmospheric N depending on site. Considering the extensive cover of shrub thickets on Virginia barrier islands, N fixation by shrubs is likely the largest single source of N to the system. The shift from grassland to shrub thicket on barrier islands results in a substantial increase in litterfall and foliar N concentration that will likely have a major impact on the size and cycling of ecosystem C and N pools. Increasing C and N availability in these nutrient-poor soils is likely to permanently reduce cover of native grasses and alter community structure by favoring species with greater N requirements.     

Temporal and structural effects of stands on litter production in Melaleuca quinquenervia dominated wetlands of south Florida

Melaleuca quinquenervia dominates large areas of the Florida Everglades in the southeastern USA where it has transformed sedge-dominated marshes into melaleuca forests. Despite its prevalence, very little is known about the ecology and stand dynamics of this invasive tree. We delineated large-, intermediate-, and small-tree stands in non-flooded, seasonally flooded and permanently flooded areas of Florida in 1997, measured their biological attributes, and then quantified litterfall components for 3–4 year periods. Melaleuca wood components and mature seed-capsules comprised the largest and the smallest portions of aboveground biomass, respectively, while leaves, fine stems, mature fruits, bud scales, floral structures, and residues represented decreasingly smaller fractions of the litter during the succeeding year. Dry weight proportion of leaves in litter was greatest (80.9%) in non-flooded and least (69.1%) in permanently flooded habitats. It was also greatest in small (85.6%) and least in large (64.7%) tree stands. Reproductive structures and mature-fruit fractions in litter were highest in large-tree stands whereas the bud-scale fraction showed no relationship to tree size. Seasonally flooded habitats had the most litterfall, wherein small-, intermediate-, and large-tree stands generated 0.662, 0.882, and 1.128 kg m⁻² yr⁻¹, respectively. Dry weight of stems, leaves, bud–scales, floral structures, and mature fruit fractions in litter increased as the predominant size of the trees in the stand increased. Total annual litter production was highest during 1999–2000. Leaf fall occurred year-round with maximal amount during April, July, and October. Highest amounts of bud scales and floral structures fell during October–January, which corresponded with flushes of vegetative growth and major flowering events. Overall, melaleuca alone accounted for nearly 99% of the total litterfall dry weight in all habitats and months sampled. The amount of non-melaleuca litter was greater in small-tree stands than in intermediate- or large-tree stands. Litterfall data of this nature will be helpful in detecting changes occurring in melaleuca canopies in response to biological control impact and in prescribing site-specific management strategies.     

Nitrogen dynamics of a boreal black spruce wildfire chronosequence

This study examined the nitrogen (N) dynamics of a black spruce (Picea mariana (Mill.) BSP)-dominated chronosequence in Manitoba, Canada. The seven sites studied each contained separate well- and poorly drained stands, originated from stand-killing wildfires, and were between 3 and 151 years old. Our goals were to (i) measure total N concentration ([N]) of all biomass components and major soil horizons; (ii) compare N content and select vegetation N cycle processes among the stands; and (iii) examine relationships between ecosystem C and N cycling for these stands. Vegetation [N] varied significantly by tissue type, species, soil drainage, and stand age; woody debris [N] increased with decay state and decreased with debris size. Soil [N] declined with horizon depth but did not vary with stand age. Total (live + dead) biomass N content ranged from 18.4 to 99.7 g N m⁻² in the well-drained stands and 37.8–154.6 g N m⁻² in the poorly drained stands. Mean soil N content (380.6 g N m⁻²) was unaffected by stand age. Annual vegetation N requirement (5.9 and 8.4 g N m⁻² yr⁻¹ in the middle-aged well- and poorly drained stands, respectively) was dominated by trees and fine roots in the well-drained stands, and bryophytes in the poorly drained stands. Fraction N retranslocated was significantly higher in deciduous than evergreen tree species, and in older than younger stands. Nitrogen use efficiency (NUE) was significantly lower in bryophytes than in trees, and in deciduous than in evergreen trees. Tree NUE increased with stand age, but overall stand NUE was roughly constant (∼ ∼150 g g⁻¹ N) across the entire chronosequence.     

Tree ring δ<Superscript>15</Superscript>N as validation of space-for-time substitution in disturbance studies of forest nitrogen status

Forest ecosystem nitrogen (N) response to disturbance has often been examined by space-for-time substitution, but there are few objective tests of the possible variation in disturbance type and intensity across chronosequence sites. We hypothesized that tree ring δ¹⁵N, as a record of ecosystem N status, could validate chronosequence assumptions and provide isotopic evidence to corroborate N trends. To test this we measured soil N availability, soil δ¹⁵N, and foliar N attributes of overstory Douglas-fir (Pseudotsuga menziesii) and understory western hemlock (Tsuga heterophylla) across three old-growth stands and nine second-growth plantations on southeast Vancouver Island, British Columbia (Canada). Increment cores for wood δ¹⁵N were retrieved from three co-dominant Douglas-fir per plot. Bulk soil δ¹⁵N was well aligned with both foliar and recent wood δ¹⁵N, demonstrating the utility of wood δ¹⁵N in monitoring ecosystem N status. Strongly contrasting trends in tree ring δ¹⁵N were evident among second-growth stands, with most trees from plantations older than 50 years exhibiting steep declines (3–4‰) in δ¹⁵N but with no temporal trends detected for younger plantations. The discrepancy in tree ring δ¹⁵N suggests disturbance history varied considerably among second-growth sites, likely because of greater slash loads and hotter broadcast burns on older cutblocks. As a consequence, the pattern of increased soil N availability and foliar N concentration with time since disturbance derived from the chronosequence could not be validated. Tree ring δ¹⁵N may provide insights into disturbance intensity, especially fire, and correlations with foliar N concentration could inform the extent of changes in stand nutrition.     

Seasonal dynamics of fine root biomass, root length density, specific root length, and soil resource availability in a Larix gmelinii plantation

Fine root turnover is a major pathway for carbon and nutrient cycling in terrestrial ecosystems and is most likely sensitive to many global change factors. Despite the importance of fine root turnover in plant C allocation and nutrient cycling dynamics and the tremendous research efforts in the past, our understanding of it remains limited. This is because the dynamics processes associated with soil resources availability are still poorly understood. Soil moisture, temperature, and available nitrogen are the most important soil characteristics that impact fine root growth and mortality at both the individual root branch and at the ecosystem level. In temperate forest ecosystems, seasonal changes of soil resource availability will alter the pattern of carbon allocation to belowground. Therefore, fine root biomass, root length density (RLD) and specific root length (SRL) vary during the growing season. Studying seasonal changes of fine root biomass, RLD, and SRL associated with soil resource availability will help us understand the mechanistic controls of carbon to fine root longevity and turnover. The objective of this study was to understand whether seasonal variations of fine root biomass, RLD and SRL were associated with soil resource availability, such as moisture, temperature, and nitrogen, and to understand how these soil components impact fine root dynamics in Larix gmelinii plantation. We used a soil coring method to obtain fine root samples (⩽2 mm in diameter) every month from May to October in 2002 from a 17-year-old L. gmelinii plantation in Maoershan Experiment Station, Northeast Forestry University, China. Seventy-two soil cores (inside diameter 60 mm; depth intervals: 0–10 cm, 10–20 cm, 20–30 cm) were sampled randomly from three replicates 25 m × 30 m plots to estimate fine root biomass (live and dead), and calculate RLD and SRL. Soil moisture, temperature, and nitrogen (ammonia and nitrates) at three depth intervals were also analyzed in these plots. Results showed that the average standing fine root biomass (live and dead) was 189.1 g·m⁻²·a⁻¹, 50% (95.4 g·m⁻²·a⁻¹) in the surface soil layer (0–10 cm), 33% (61.5 g·m⁻²·a⁻¹), 17% (32.2 g·m⁻²·a⁻¹) in the middle (10–20 cm) and deep layer (20–30cm), respectively. Live and dead fine root biomass was the highest from May to July and in September, but lower in August and October. The live fine root biomass decreased and dead biomass increased during the growing season. Mean RLD (7,411.56 m·m⁻³·a⁻¹) and SRL (10.83 m·g⁻¹·a⁻¹) in the surface layer were higher than RLD (1 474.68 m·m⁻³·a⁻¹) and SRL (8.56 m·g⁻¹·a⁻¹) in the deep soil layer. RLD and SRL in May were the highest (10 621.45 m·m⁻³ and 14.83m·g⁻¹) compared with those in the other months, and RLD was the lowest in September (2 198.20 m·m⁻³) and SRL in October (3.77 m·g⁻¹). Seasonal dynamics of fine root biomass, RLD, and SRL showed a close relationship with changes in soil moisture, temperature, and nitrogen availability. To a lesser extent, the temperature could be determined by regression analysis. Fine roots in the upper soil layer have a function of absorbing moisture and nutrients, while the main function of deeper soil may be moisture uptake rather than nutrient acquisition. Therefore, carbon allocation to roots in the upper soil layer and deeper soil layer was different. Multiple regression analysis showed that variation in soil resource availability could explain 71–73% of the seasonal variation of RLD and SRL and 58% of the variation in fine root biomass. These results suggested a greater metabolic activity of fine roots living in soil with higher resource availability, which resulted in an increased allocation of carbohydrate to these roots, but a lower allocation of carbohydrate to those in soil with lower resource availability. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(3): 403–410 [译自: 植物生态学报, 2005, 29(3): 403–410]     

Carbon, nitrogen and phosphorus in volcanic soils following afforestation with native birch (Betula pubescens) and introduced larch (Larix sibirica) in Iceland

Afforestation has become an important tool for soil protection and land reclamation in Iceland. Nevertheless, the harsh climate and degraded soils are growth-limiting for trees, and little is know about changes in soil nutrients in maturing forests planted on the volcanic soils. In the present chronosequence study, changes in C, N and total P in soil (0–10 and 10–20 cm depth) and C and N in foliar tissue were investigated in stands of native Downy birch (Betula pubescens Enrh.) and the in Iceland introduced Siberian larch (Larix sibirica Ledeb.). The forest stands were between 14 and 97 years old and were established on heath land that had been treeless for centuries. Soils were Andosols derived from basaltic material and rhyolitic volcanic ash. A significant effect of tree species was only found for the N content in foliar tissue. Foliar N concentrations were significantly higher and foliar C/N ratios significantly lower in larch needles than in birch leaves. There was no effect of stand age. Changes in soil C and the soil nutrient status with time after afforestation were little significant. Soil C concentrations in 0–10 cm depth in forest stands older than 30 years were significantly higher than in heath land and forest stands younger than 30 years. This was attributed to a slow accumulation of organic matter. Soil N concentrations and soil P_tot were not affected by stand age. Nutrient pools in the two soil layers were calculated for an average weight of soil material (400 Mg soil ha⁻¹ in 0–10 cm depth and 600 Mg soil ha⁻¹ in 10–20 cm depth, respectively). Soil nutrient pools did not change significantly with time. Soil C pools were in average 23.6 Mg ha⁻¹ in the upper soil layer and 16.9 Mg ha⁻¹ in the lower soil layer. The highest annual increase in soil C under forest compared to heath land was 0.23 Mg C ha⁻¹ year⁻¹ in 0–10 cm depth calculated for the 53-year-old larch stand. Soil N pools were in average 1.0 Mg N ha⁻¹ in both soil layers and did not decrease with time despite a low N deposition and the uptake and accumulation of N in biomass of the growing trees. Soil P_tot pools were in average 220 and 320 kg P ha⁻¹ in the upper and lower soil layer, respectively. It was assumed that mycorrhizal fungi present in the stands had an influence on the availability of N and P to the trees. Responsible Editor: Hans Lambers.     

Fine root dynamics in lodgepole pine and white spruce stands along productivity gradients in reclaimed oil sands sites

Open‐pit mining activities in the oil sands region of Alberta, Canada, create disturbed lands that, by law, must be reclaimed to a land capability equivalent to that existed before the disturbance. Re‐establishment of forest cover will be affected by the production and turnover rate of fine roots. However, the relationship between fine root dynamics and tree growth has not been studied in reclaimed oil sands sites. Fine root properties (root length density, mean surface area, total root biomass, and rates of root production, turnover, and decomposition) were assessed from May to October 2011 and 2012 using sequential coring and ingrowth core methods in lodgepole pine (Pinus contorta Dougl.) and white spruce (Picea glauca (Moench.) Voss) stands. The pine and spruce stands were planted on peat mineral soil mix placed over tailings sand and overburden substrates, respectively, in reclaimed oil sands sites in Alberta. We selected stands that form a productivity gradient (low, medium, and high productivities) of each tree species based on differences in tree height and diameter at breast height (DBH) increments. In lodgepole pine stands, fine root length density and fine root production, and turnover rates were in the order of high > medium > low productivity sites and were positively correlated with tree height and DBH and negatively correlated with soil salinity (P < 0.05). In white spruce stands, fine root surface area was the only parameter that increased along the productivity gradient and was negatively correlated with soil compaction. In conclusion, fine root dynamics along the stand productivity gradients were closely linked to stand productivity and were affected by limiting soil properties related to the specific substrate used for reconstructing the reclaimed soil. Understanding the impact of soil properties on fine root dynamics and overall stand productivity will help improve land reclamation outcomes.     

Nutrient resorption responses to water and nitrogen amendment in semi-arid grassland of Inner Mongolia, China

Litterfall and fine root production is a major pathway for carbon and nutrient cycling in forest ecosystems. We investigated leaf litterfall, fine-root mass, production and turnover rate in the upper soil (0–30 cm) under four major tree species (Leucaena leucocephala, Acacia nilotica, Azadirachta indica, Prosopis juliflora) of the semi-arid region of India. All the four tree species showed an unimodal peak of leaf litterfall with distinct seasonality. Leucaena leucocephala and Acacia nilotica had maximum leaf litterfall between September and December while Azadirachta indica and Prosopis juliflora shed most of their leaves between February and May. Annual leaf litterfall of the four species ranged from 3.3 Mg ha^?1 (Leucaena leucocephala) to 8.1 Mg ha^?1 (Prosopis juliflora). Marked seasonal variations in amount of fine root biomass were observed in all the four tree species. Fine root production was maximum in Prosopis juliflora (171 g m^?2 y^?1) followed by Azadirachta indica (169 g m^?2 y^?1), Acacia nilotica (106 g m^?2 y^?1) and Leucaena leucocephala (79 g m^?2 y^?1). Fine root biomass showed a seasonal peak after the rainy season but fell to its lowest value during the winter and dry summer season. Fine root turnover rate ranged from 0.56 to 0.97 y^?1 and followed the order Azadirachta indica > Leucaena leucocephala > Prosopis juliflora > Acacia nilotica. The results of this study demonstrated that Prosopis juliflora and Azadirachta indica had greater capability for maintaining site productivity as evidenced from greater leaf litterfall and fine root production.