Greater accumulation of litter in spruce (Picea abies) compared to beech (Fagus sylvatica) stands is not a consequence of the inherent recalcitrance of needles

Background and aims

Replacement of beech by spruce is associated with changes in soil acidity, soil structure and humus form, which are commonly ascribed to the recalcitrance of spruce needles. It is of practical relevance to know how much beech must be admixed to pure spruce stands in order to increase litter decomposition and associated nutrient cycling. We addressed the impact of tree species mixture within forest stands and within litter on mass loss and nutritional release from litter.

Methods

Litter decomposition was measured in three adjacent stands of pure spruce (Picea abies), mixed beech-spruce and pure beech (Fagus sylvatica) on three nutrient-rich sites and three nutrient-poor sites over a three-year period using the litterbag method (single species and mixed species bags).

Results

Mass loss of beech litter was not higher than mass loss of spruce litter. Mass loss and nutrient release were not affected by litter mixing. Litter decay indicated non-additive patterns, since similar remaining masses under pure beech (47%) and mixed beech-spruce (48%) were significantly lower than under pure spruce stands (67%). Release of the main components of the organic substance (C_org, N_tot, P, S, lignin) and associated K were related to mass loss, while release of other nutrients was not related to mass loss.

Conclusions

In contradiction to the widely held assumption of slow decomposition of spruce needles, we conclude that accumulation of litter in spruce stands is not caused by recalcitrance of spruce needles to decay; rather adverse environmental conditions in spruce stands retard decomposition. Mixed beech-spruce stands appear to be as effective as pure beech stands in counteracting these adverse conditions.     

模拟N沉降下不同林龄马尾松林凋落叶分解-土壤C、N化学计量特征

模拟N沉降对森林生态系统的影响是当今全球变化生态学研究的一个热点问题,土壤碳库对N沉降比较敏感,N沉降增加了凋落叶分解过程中外源N含量,间接影响凋落叶分解的化学过程并改变凋落叶分解速率,因此,研究模拟N沉降下凋落叶分解-土壤C-N关系对预测森林C吸存有重要意义。利用原位分解袋法研究了模拟N沉降下三峡库区不同林龄马尾松林(Pinus massoniana)凋落叶分解过程中凋落叶-土壤C、N化学计量响应及其关系;N沉降水平分对照(CK,0 g m~(-2)a~(-1))、低氮(LN,5 g m~(-2)a~(-1))、中氮(MN,10 g m~(-2)a~(-1))和高氮(HN,15 g m~(-2)a~(-1))。结果表明:分解540 d后,N沉降促进20年生和30年生马尾松林凋落叶分解,46年生马尾松林中仅低氮处理促进凋落叶分解,4种处理均是30年生分解最快,说明同一树种起始N含量低的凋落叶对N沉降呈正响应,N沉降处理促进起始N含量低的凋落叶分解,起始N含量高的凋落叶分解过程中易达到"N饱和"。N沉降抑制20年生和46年生凋落叶C释放(低于对照0.62%—6.69%),促进30年生C释放(高于对照0.28%—5.55%);30年生和46年生林分N固持量均高于对照(高于对照0.15%—21.34%),20年生则低于对照(5.70%—13.87%),说明模拟N沉降处理促进起始C含量低的凋落叶C释放和起始N含量低的凋落叶N固持。N沉降处理下仅30年生马尾松林土壤有机碳较对照增加,且土壤有机质与凋落叶C、N和分解速率呈正相关,与凋落叶C/N比呈显著负相关;土壤总氮与凋落叶分解速率、凋落叶N含量呈正相关,土壤有机碳/总氮比与凋落叶C、N含量呈正相关;对照处理中凋落叶分解指标对土壤养分影响顺序是分解速率凋落物C含量凋落物C/N比凋落物N含量,低、中、高氮处理中则是凋落物C含量分解速率凋落物N含量凋落物C/N比。研究表明低土壤养分含量马尾松林对N沉降呈正响应,N沉降促进低土壤养分马尾松林凋落叶分解并提高土壤肥力;凋落叶质量和土壤养分含量低的生态系统土壤C对N沉降响应更显著。     

Nutrient return with leaf litter fall in Fagus sylvatica forests across a soil fertility gradient

Leaf litter fall is an important nutrient flux in temperature deciduous forests which supplies a large part of the rapidly mineralisable nutrient fraction to the soil. This study investigates nutrient return with leaf litter fall in 36 old-growth forest stands of Fagus sylvatica across a broad gradient of soil fertility covering 9 mesozoic and kaenozoic parent material types (three limestones, two sandstones, two clay stones, one sand and one loess substrate). Study objectives were to analyse (i) the dependency of leaf litter nutrient concentrations on soil fertility, and (ii) the relationship between soil fertility and nutrient return with leaf litter at the stand level. Beech stands on the nine parent material types produced similar annual leaf litter masses irrespective of soil fertility or acidity. Leaf litter from the nine parent materials showed only minor variation with respect to N and K concentrations (factors of 1.5 and 1.4), moderate variation for Ca, Mg and P concentrations (factors of 2.2 to 2.9), and high variation for Al and Mn concentrations (factors of 6.7 and 10.5). Consequently, annual nutrient return with litter fall (leaf litter mass x litter nutrient concentration) was more similar among the parent materials for N (165–273 mmol m⁻² yr1) ⁻¹ and K (16–30 mm m⁻² yr⁻¹) than for Ca, P, Mg, Mn and Al. A possible explanation is increased N deposition in recent time. According to a correlation analysis, return rates of N, P, K and Mg (but not Ca) were independent of the pool size of the respective nutrient in the soil. N return rate was neither influenced by the soil pools of Nt, plant- available P (Pa) or exchangeable Ca, K and Mg, nor by soil acidity or the exchangeable Al pool. P return, in contrast, showed a negative relation to soil fertility. We hypothesize that nutrient fluxes with leaf litter fall do not necessarily reduce the fitness of tree populations as has been postulated from a tree-centred view. Rather, we suggest that nutrient fluxes with litter fall can increase, instead of decrease, plant fitness by improving nutrient availability in the densely rooted topsoil which reduces the roots’ carbon and nutrient costs of nutrient acquisition.     

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.     

Plant-soil feedback in spruce (Picea abies) and mixed spruce-beech (Fagus sylvatica) stands as indicated by dendrochemistry

Ten pairs of secondary pure spruce (Picea abies) and adjacent mixed spruce-beech (Fagus sylvatica) stands on comparable sites were selected on two different bedrocks for soil formation (Flysch: nutrient rich and high soil pH; Molasse: poor nutrient supply and acidic) to study how an admixture of beech to spruce stands affects nutrient cycling and consequently soil chemistry. Soil analyses indicated accumulation of Ca under the mixed stands while the top soil under pure spruce was acidified. It was hypothesized that changes of soil chemical properties due to species composition over the last six decades are reflected in the stem wood of spruce. Three healthy dominant spruce trees per plot were selected for coring. Cores were crossdated and half-decadal samples were analyzed for Ca, Mg, Mn and Al. Calcium and Mg concentrations in stem wood of spruce were significantly higher for the pure spruce than for the mixed stands in spite of lower Ca and Mg stores in the soil. We assume that acidification caused by pure spruce mobilized these cations temporarily, increasing soil solution contents and consequently stem wood concentrations. It was possible to reconstruct soil pH from the element ratios Ca/Al (pure stands) and Ca/Mg (mixed stands), since these ratios in the stem wood of the last half-decade did correlate with soil pH for selected soil depths. Reconstructed soil pH showed a decline over the last 60 years under both species compositions due to accumulation of base cations in the increasing biomass. Comparisons of reconstructed soil pH in 0–5 and 10–20 cm soil depth indicated more pronounced top soil acidification (lower soil pH in 0–5 cm) by spruce on the nutrient rich soil (Flysch) than on the acidic soil (Molasse). However, admixture of beech caused higher pH values in 0–5 cm than in 10–20 cm soil depth on Flysch due to the observed Ca-pump effect of beech (uptake of Ca from deeper soil horizons).     

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.     

Leguminous Trees for the Rehabilitation of Sodic Wasteland in Northern India

The study objective was to determine the fuelwood harvesting rotation and magnitude of nutrient accumulation by the sodic tolerant species Dalbergia sissoo and Prosopis juliflora for the rehabilitation of sodic wastelands. Mean annual increment and current annual increment growth of these trees reached a peak in six-year-old stands. Leaves dominated the litter of these species and contained high concentrations of most nutrients. N, Ca, Mg, and Fe were present in larger concentrations than other nutrients. Organic carbon additions at harvest after six years averaged 16 Mg ha ^{? 1}, while soil pH was reduced from 9.8 to 8.6. Exchangeable sodium percentage (ESP) values decreased 65 and 29% under P. juliflora and D. sissoo, respectively, after harvest. These tree species produced significant root spread and deep penetration and were able to rehabilitate sodic soil effectively.     

Coupled Nitrogen and Calcium Cycles in Forests of the Oregon Coast Range

Nitrogen (N) is a critical limiting nutrient that regulates plant productivity and the cycling of other essential elements in forests. We measured foliar and soil nutrients in 22 young Douglas-fir stands in the Oregon Coast Range to examine patterns of nutrient availability across a gradient of N-poor to N-rich soils. N in surface mineral soil ranged from 0.15 to 1.05% N, and was positively related to a doubling of foliar N across sites. Foliar N in half of the sites exceeded 1.4% N, which is considered above the threshold of N-limitation in coastal Oregon Douglas-fir. Available nitrate increased five-fold across this gradient, whereas exchangeable magnesium (Mg) and calcium (Ca) in soils declined, suggesting that nitrate leaching influences base cation availability more than soil parent material across our sites. Natural abundance strontium isotopes (⁸⁷Sr/⁸⁶Sr) of a single site indicated that 97% of available base cations can originate from atmospheric inputs of marine aerosols, with negligible contributions from weathering. Low annual inputs of Ca relative to Douglas-fir growth requirements may explain why foliar Ca concentrations are highly sensitive to variations in soil Ca across our sites. Natural abundance calcium isotopes (δ⁴⁴Ca) in exchangeable and acid leachable pools of surface soil measured at a single site showed 1 per mil depletion relative to deep soil, suggesting strong Ca recycling to meet tree demands. Overall, the biogeochemical response of these Douglas-fir forests to gradients in soil N is similar to changes associated with chronic N deposition in more polluted temperate regions, and raises the possibility that Ca may be deficient on excessively N-rich sites. We conclude that wide gradients in soil N can drive non-linear changes in base-cation biogeochemistry, particularly as forests cross a threshold from N-limitation to N-saturation. The most acute changes may occur in forests where base cations are derived principally from atmospheric inputs.     

Cation distribution,cycling, and removal from mineral soil in Douglas-fir and red alder forests

Overstory species influence the distribution and dynamics of nutrients in forest ecosystems. Ecosystem-level estimates of Ca, Mg, and K pools and cycles in 50-year old Douglas-fir and red alder stands were used to determine the effect of overstory composition on net cation removal from the mineral soil, i.e. cation export from the soil in excess of additions. Net cation removal from Douglas-fir soil was 8 kg Ca ha^–1 yr^–1, 1 kg Mg ha^–1 yr^–1, and 0.3 kg K ha^–1 yr^–1. Annual cation export from soil by uptake and accumulation in live woody tissue and O horizon was of similar magnitude to leaching in soil solution. Atmospheric deposition partially off-set export by adding cations equivalent to 28–88% of cation export. Net cation removal from red alder soil was 58 kg Ca ha^–1 yr^–1, 9 kg Mg ha^–1 yr^–1, and 11 kg K ha^–1 yr^–1. Annual cation accumulation in live woody tissue and O horizon was three times greater than in Douglas-fir, while cation leaching in soil solution was five to eight times greater. The lack of excessive depletion of exchangeable cations in the red alder soil suggests that mineral weathering, rather than exchangeable cations, was the source of most of the removed cations. Nitric acid generated during nitrification in red alder soil led to high rates of weathering and NO₃-driven cation leaching.     

Effects of forest clear-cutting on the sulphur, phosphorus and base cations fluxes through podzolic soil horizons

Clear-cutting considerably alters the flow of nutrients through the forest ecosystem. These changes are reflected in soil solution concentrations and fluxes. The effects of clear-cutting (stems only) on the fluxes of water soluble phosphorus (P), sulphur (S) and base cations (Ca, Mg and K) through a podzolic soil were studied in a Norway spruce dominated mixed boreal forest in eastern Finland. Bulk deposition, total throughfall (throughfall + stemflow) and soil percolate from below the organic (O), eluvial (E) and illuvial (B) horizons were collected for 4 years before and for 3 years after cutting. Annual deposition loads (kg ha^–1) to the forest floor were less after clear-cutting, averaging 1.7 S, 0.84 Ca, 0.14 Mg, 0.64 K and 0.10 P. Before cutting, the loads were 4.6 S, 2.7 Ca, 0.70 Mg, 6.2 K and 0.20 P. Annual fluxes of total S and sulphate (SO ₄ ^2– ) from below the O-horizon were also lower (33%) after clear-cutting, total S averaging 2.0 kg ha^–1, the flux from below the B-horizon also diminished after clear-cutting. The flux of total P (mainly inorganic) from below the O-horizon increased threefold (6.9 kg ha^–1; sum over the 3-year period) compared to period before cutting. The fluxes of base cations from below the O-horizon increased twofold. The flux of K⁺ from below the O- and E-horizons was most strongly correlated with that of phosphate (PO ₄ ^3– ) and those of Ca²⁺ and Mg²⁺ with the DOC flux. Increased fluxes of P and base cations to the mineral soil generated only slightly increased fluxes from below the B-horizon. The retention of base cations and P in the mineral soil indicates there was little change in leaching to ground and surface waters after clear-cutting.     

滇中常绿阔叶林及云南松林水文作用的初步研究

 本文研究了滇中地区山地常绿阔叶林及云南松林的林冠截留量,冠流和茎流量,地表水土流失量,林地枯枝落叶层持水量,土壤含水量以及雨水、冠流、茎流和地表迳流中营养元素(N、P、K、Ca和Mg)的含量,并分析比较了针、阔两类森林调节和涵养水分的作用和降水中林冠养分的淋溶、迁移特点,对了解滇中亚热带山地森林生态系统的功能和生产力的研究,对本区森林资源的保护,林业的合理经营,均有重要意义。     

不同林龄马尾松凋落物基质质量与土壤养分的关系

凋落物的质量、数量及分解速率在一定程度上代表了土壤的营养状况。为了精确估算凋落物分解对土壤碳库的年净归还量及凋落物-土壤生物化学连续体的深层理解,从凋落物基质质量的角度分析了三峡库区不同林龄马尾松凋落物基质质量与土壤养分的作用关系,结果表明:中龄林、近熟林、成熟林马尾松凋落物基质质量中的C、C/N比、C/P比、木质素/N比、木质素/P比差异显著,其中近熟林凋落物叶木质素/N分别比中龄林和成熟林的高33.65%、39.24%,N、P、K、木质素含量差异不显著;但各组织器官的N、P、K含量差异显著,均是皮<枝<叶<杂物,C/N比、C/P比的变化则相反。不同林龄马尾松0-20 cm(0-5 cm、5-10 cm、10-20 cm)土壤有机质、总氮、有效磷含量均表现出近熟林<中龄林<成熟林,0-5 cm最大,10-20 cm最小,且随着土壤深度的增加而明显降低,总磷则是中林龄最低,成熟林最大,pH值则各土层均表现为中龄林<成熟林<近熟林,平均pH值为4.55-5.51。凋落物基质质量指标与土壤养分之间冗余分析(RDA)表明:马尾松凋落物基质质量和土壤养分之间关系紧密,N、P、纤维素、半纤维素、木质素、木质素/N比、C/N比对土壤养分影响比较大;凋落物中木质素/N比、C/N比与土壤有机质呈显著负相关,其含量越高越不利于土壤有机质的形成,土壤养分积累的越慢;凋落物基质质量氮含量与土壤氮含量呈显著正相关;土壤pH值、容重与N含量呈显著负相关,与凋落物C/N比、木质素/N比呈显著正相关。马尾松土壤表面有机质、N、P养分含量与凋落物基质质量对应养分含量变化规律一致,土壤养分高,凋落物基质质量相对较高,土壤贫瘠,凋落物基质质量相对较低。     

Deterioration of Norway spruce vitality despite a sharp decline in acid deposition: a long‐term integrated perspective

Since the late 1970s, several long‐term ecological studies were conducted to better understand the biogeochemical functioning of Norway spruce stands in the Ardennes as these nutrient‐poor ecosystems were subject to high levels of acid deposition and exhibited symptoms of tree health decline. Between 1978 and 2009, acid deposition declined sharply, especially sulfur and to a lesser extent nitrogen deposition. The aim of this study was (i) to determine if the Norway spruce stands recovered after the reduction of acid deposition and (ii) to explain why such a recovery occurred or not. Therefore, we collected data from different projects carried out in the Ardennes to characterize the long‐term temporal trends in soil solution chemistry, foliar nutrition, and crown condition. In parallel, a model describing the nutrient cycling in forests (NuCM) was calibrated and used to check the consistency of the observed temporal trends and to explain them. The soil solution concentration of most of the elements decreased between 1978 and 2002, which was ascribed to a decrease in atmospheric deposition. For potassium, a decline in the exchangeable pool was also showed based on the simulation carried out with NuCM. As nitrogen (N) deposition remained at an elevated level, Norway spruce stands were progressively saturated in N and mineral nutrition became more and more unbalanced. Except the foliar N and Al concentration that remained constant and increased respectively, the foliar concentration of all other nutrients decreased between 1993 and 2009, which can be explained by the decrease in ion concentration in solution. These nutritional disorders weakened trees and were probably exacerbated during the 2003 summer drought, after which symptoms of vitality loss progressively appeared. In these N‐saturated ecosystems, the N cycle was disrupted by this health decline, which increased NO₃^? leaching reinforcing soil acidification and risk of aluminum (Al) toxicity.     

Nutrients and mass in litter and top soil of ten tropical tree plantations

The importance of litter to nutrient and organic matter storage and the possible influence of species selection on soil fertility in ten stands each consisting of a separate tree species were examined in this study. The plantations had been grown under similar conditions in an arboretum in the Luquillo Experimental Forest, Puerto Rico. The species involved were: Anthocephalus chinensis, Eucalyptus × patentinervis, E. saligna, Hernandia sonora, Hibiscus elatus, Khaya nyasica, Pinus caribaea var. hondurensis, P. elliottii var. densa, Swietenia macrophylla, and Terminalia ivorensis. After 26 yr, litter mass ranged from 5 mg ha^-1 in the H. sonora stand to 27.2 Mg ha^-1 in the P. caribaea stand. Nutrients in the litter (N, P, K, Ca, and Mg) also varied widely, but stands were ranked in different order when ranked by nutrients in the litter than then ranked according to accumulation of mass. Only E. saligna and A. chinensis stands were ranked similarly in accumulation of both nutrients and mass, and the stand of H. elatus was ranked higher with respect to nutrient accumulation than to accumulation of mass. The nutrient concentration in standing leaf litter generally increased in the order of recently fallen <old intact< fragmented. Nutrient concentration of standing leaf litter appears to increase with age and depth in the litter layer. The amount of nutrients stored in the litter compartment of these plantations was in the same order of magnitude as the quantity of available nutrients in the top 10-cm of mineral soil. Total litter mass was negatively correlated with the mass-weighted concentration of N, K, and Mg. The same relationship was found for Ca in the leaf litter and N in the fine wood litter compartments. In some stands (notably P. caribaea, P. elliottii, and E. saligna), leaf litter derived from species other than the species planted in that particular stand had higher nutrient concentration than leaf litter from the planted species. Soils of the 10 stands were classified in the same soil series and had similar texture (clay soils). However, significantly different chemical characteristics were found. Results obtained by analysis of covariance and by limiting comparisons to adjacent stands with similar soil texture, indicate that different species have had different influences on the concentration of available nutrients in soil.     

加勒比松林凋落物对地表径流和氮、磷流失的影响

以佛山的加勒比松林(Pinus caribaea)为对象, 研究了其地表径流及N和P流失量特点。结果表明, 有凋落物加勒比松林和无凋落物加勒比松林的年径流量分别为12.6和51.8 mm, 其中夏季占90%以上。2种林分的地表径流量和降雨量的关系遵循线性方程。除2007年7月和2008年2月外, 各月地表径流N浓度为有凋落物加勒比松林大于无凋落物加勒比松林, 浓度的差异主要是因为年径流量不同引起的。总体来看, 有凋落物加勒比松林的地表径流P浓度高, 无凋落物加勒比松林低。地表径流N浓度与径流量之间存在着负指数关系。有凋落物加勒比松林和无凋落物加勒比松林的地表径流总N输出量分别为341和983 g·hm^-2, 夏季分别占其年N流失量的73%和64%。无凋落物加勒比松林的地表径流P流失量高达21.5 g·hm^-2, 而有凋落物加勒比松林为6.5 g·hm^-2。2008年6月为有凋落物加勒比松林和无凋落物加勒比松林的地表径流P流失高峰, 分别占其年流失量的46%和60%。因此, 保留林下凋落物可以降低加勒比松林的N和P的流失。     

Nutrient cycling in an excessively rainfed subtropical grassland at Cherrapunji

Cycling of six mineral elements (N, P, K, Na, Ca and Mg) was studied in a humid subtropical grassland at Cherrapunji, north-eastern India during 1988-1989. Elemental concentrations in the shoot of four dominant grass species,viz., Arundinella khaseana, Chrysopogon gryllus, Eragrostiella leioptera andEulalia trispicata were very low, and none of the species appears suitable for fodder use. Among different vegetation compartments, live root was the largest reservoir of all the nutrients (except Ca) followed by live shoot, dead shoot, litter and dead root. For Ca, live shoot was the major storage compartment. The total annual uptake (kg ha^-1) was 137.3, 10.4, 51.1, 5.5, 8.7 and 18.2 for N, P, K, Na, Ca and Mg, respectively. In an annual cycle 98% N, 77% P, 49% K, 109% Na, 87% Ca and 65% Mg returned to the soil through litter and belowground detritus. A major portion of N, P and Na was recycled through the belowground system, whereas nearly half of K, Ca and Mg was recycled through the shoot system. Precipitation acts as the source of N and P input, but at the same time causes loss of cations.     

Effects of litter removal on arthropod communities in pine plantations

Natural and anthropogenic disturbances can cause abrupt changes in trophic interactions by altering the rate, timing, or composition of organic inputs to ecological systems which in turn can shift patterns of species dominance. We examined the short-term effects of litter removal on soil fauna in pine plantations of three different species (longleaf, Pinus palustris; loblolly, P. taeda; and slash, P. elliottii) using a manipulative experiment, with the goal of examining differences among dominant orders of arthropods and differences among timber types. We sampled arthropods once per month for 6 months immediately following raking, and found that removal of the litter caused significant changes to abundance or presence of five of the nine dominant orders. Reductions in abundances of arthropod orders were most apparent in loblolly pine stands, while increases in abundance were more common in longleaf and slash pine stands. The differential impact among orders suggests that removal of the litter layer is likely to alter trophic interactions by changing the relative abundance of functional groups. Repeated litter removal via raking could have negative repercussions on ecosystem stability. Finally, nutrient additions through fertilization seem unlikely to mitigate the changes imposed on the arthropod community through litter removal.     

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.     

Impact of common European tree species and Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) on the physicochemical properties of the rhizosphere

Trees play a crucial role in nutrient cycling and ecosystem fertility, notably through rhizosphere processes. The aim of this study was to compare soil physicochemical properties between bulk soil and rhizosphere of several tree species, and to compare rhizosphere properties between fertilized and non-fertilized conditions. The soil sampling was performed in Breuil-Chenue forest (North-East of France) in seven stands: native forest (old beech (Fagus sylvatica L.) and oak (Quercus sessiliflora Smith) coppice with standards; CwS), beech, oak (Quercus petraea [Matt.] Liebl.), Douglas-fir and fertilised Douglas-fir, Norway spruce (Picea abies Karst.) and fertilised Norway spruce. Systematic soil sampling was performed at 0–3, 3–10, and 10–23 cm in 20 calibrated pits. The rhizosphere of the different species was generally enriched in C, N, Ca, Mg, and K. Interestingly, the same positive effect was observed in the fertilised plots. The rhizosphere effect varied between tree species for C, “base” cations, pH_water and cation exchange capacity. This study reveals that interactions between roots, microorganisms and soil can enrich the pool of nutrients in the rhizosphere compared to bulk soil whatever the soil fertility conditions, and that the magnitude of the rhizosphere effect depends on tree species.     

天然枫桦红松林凋落量动态及养分归还量

三年定位研究表明小兴安岭天然枫桦红松林年均凋落量有5.8t／ha(干重)。凋落量的季节变化格局是随着气候变冷有一明显秋季凋落高峰期(9—10月)。凋落物每年养分的归还量：Ca、N、K、Mg、P,相应为67.0、56.9、14.8、9.5、和6.6kg／ha,总计155.0kg／ha。据测定阔叶树落叶养分含量明显高于所有针叶的含量。尽管阔叶树的年凋落量仅占该混交林的年总凋落量的三分之一,阔叶树落叶仍有相当高的养分比例(43.4%)归还土壤。因此,红松林分的经营管理中,保留适当比重的阔叶树有利于土壤改良和促进林分生长。