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

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

Spatial variation and understorey competition effect of Pinus radiata fine roots in a silvopastoral system in New Zealand

In designing agroforestry systems, the combination of tree genotype (orspecies) and pasture species and the spatial arrangement of trees are importantconsiderations. The spatial variation of fine root length density (FRLD) ofthree radiata pine (Pinus radiata D. Don) genotypes,referred to here as clone 3, clone 4 and seedlings, was studied in athree-year-old temperate silvopastoral experiment. The genotypes were plantedwith three understorey types: ryegrass (Lolium perenne)mixed with clovers (Trifolium spp), lucerne(Medicago sativa), and control (bare ground). Also fineroot distribution of both tree and pasture species with soil depth and inrelation to tree row (0.9 m north or south of and within the rippedtree row) was studied. Greater FRLD was found in clonal than in seedling treesin the bare ground treatment but not in the two pasture treatments, and in the0–0.1 m but not in the 0.1–0.2 or 0.2–0.3m soil layers. Clonal trees had a greater ability to develop a moreextensive root system, especially in the 0–0.1 m soil layer,but that advantage disappeared when they were planted with pasture species sincecompetition from the pasture species was most severe in the 0–10cm layer. The FRLD of lucerne was greater than that ofryegrass/clovers, consistent with the greater aboveground biomass production oflucerne. Pasture species FRLD was greater on the south (wetter) than on thenorth side of the ripline or in the ripline. The interception of prevailingsoutherly rain-bearing wind by tree crowns resulted in the south side beingwetter than the north side. Results indicated that production and distributionof fine roots of both tree and pasture species responded to changes in themicroclimate. We suggest that to optimize pasture/tree biomass productionplanting trees in the north-south direction is better than in the east-westdirection at the studied site. This revised version was published online in June 2006 with corrections to the Cover Date.     

徐州云龙山侧柏林细根形态与分布规律研究

为了提高徐州市石灰岩山地侧柏人工林的生态和景观质量,该研究以云龙山侧柏人工林为对象,利用Win-RHIZO根系分析仪对2个样地不同土层深度细根的形态参数、比根长、根长密度和生物量进行了综合分析。结果表明：（1）细根直径随根序等级的升高而增加,细根长度、比根长、表面积表现为下降趋势,而细根体积表现为先减小后增加的趋势;（2）在不同立地条件下,细根长度、比根长、表面积和体积均表现为表层高于底层,东坡高于西坡的规律;（3）单位面积上细根的生物量随根序的升高而增加,而根长密度则明显下降;表土层中细根生物量和根长密度都要高于底层,东坡高于西坡。     

Soil respiration and microbial biomass in a pecan — cotton alley cropping system in Southern USA

Little information is available on soil respiration and microbial biomass in soils under agroforestry systems. We measured soil respiration rate and microbial biomass under two age classes (young and old) of a pecan (Carya illinoinensis) — cotton (Gossypium hirsutum) alley cropping system, two age classes of pecan orchards, and a cotton monoculture on a well-drained, Redbay sandy loam (a fine-loamy, siliceous, thermic Rhodic Paleudult) in southern USA. Soil respiration was quantified monthly during the growing season from May to November 2001 using the soda-lime technique and was corrected based on infrared gas analyzer (IRGA) measurements. The overall soil respiration rates ranged from 177 to 776 mg CO₂ m^–2 h^–1. During the growing season, soil respiration was higher in the old alley cropping system than in the young alley cropping system, the old pecan orchard, the young pecan orchard, and the monoculture. Microbial biomass C was higher in the old alley cropping system (375 mg C kg^–1) and in the old pecan orchard (376 mg C kg^–1) compared to the young alley cropping system (118 mg C kg^–1), young pecan orchard (88 mg C kg^–1), and the cotton monoculture (163 mg C kg^–1). Soil respiration was correlated positively with soil temperature, microbial biomass, organic matter, and fine root biomass. The effect of alley cropping on soil properties during the brief history of alley cropping was not significant except in the old systems, where there was a trend of increasing soil respiration with short-term alley cropping. Over time, different land use and management practices influenced soil properties such as soil temperature, moisture, microbial biomass, organic matter, and fine root biomass, which in turn affected the magnitude of soil respiration. Our results suggest that trees in agroforestry systems have the potential to enhance soil fertility and sustainability of farmlands by improving soil microbial activity and accreting residual soil carbon.This revised version was published online in November 2005 with corrections to the Cover Date.     

渭北主要造林树种细根生长及分布与土壤密度关系

采用野外样地调查和盆栽试验的方法 ,研究了土壤密度对渭北主要造林树种细根生长及分布特征的影响。结果表明 ,在 0～ 10 0cm的深度内 ,不论是阴坡还是阳坡 ,油松林地的土壤密度均表现出随土层的加深逐渐增大的趋势 ;刺槐林地的土壤密度大于油松林地 ,且各土层间的差异不大。油松林地各土层土壤密度与不同径级根系长度存在显著的负相关关系。盆栽基质密度对不同树种苗木根系的生长和发育的影响虽各异 ,但在基质密度和孔隙度适中的条件下均生长良好 ,生物量达到最大。盆栽基质的密度对刺槐和油松根系的生长和发育的影响最显著 ;侧柏和山杏的细根生长发育受土壤密度的影响较小 ,其根系生物量在不同密度的基质中没有明显变化。     

Estimating the production and mortality of fine roots in a Japanese cedar (Cryptomeria japonica D. Don) plantation using a minirhizotron technique

Fine roots are a key component of forested ecosystems, but available information is still limited. This study examined the production and mortality of fine roots less than 1 mm in diameter in a Japanese cedar (Cryptomeria japonica D. Don) plantation located on the Kanto Plain in central Japan. We used a minirhizotron technique in combination with soil coring, and collected data for 1 year (May 2002–May 2003). Fine root production and mortality were determined from changes in the lengths of individual fine roots on minirhizotron tubes. Both fine root production and mortality rates were greater in the upper soil than in lower soil levels. Both rates were seasonal, with higher values in summer than in winter; this trend was more pronounced in upper soil levels. These results suggest that environmental conditions, such as temperature or soil properties, affect the production and mortality rates of fine roots. Fine root production and mortality occurred simultaneously, and their rates were similar, which may have led to unclear seasonal changes in fine root standing crop estimates. Soil coring indicated that the fine root biomass of this stand was about 120 g m⁻², of which 40% was from Japanese cedar. The estimated rates of dry matter production and mortality of total fine roots, including understory plants, were both approximately 300 g m⁻² year⁻¹.     

Root development across a chronosequence in a Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don) plantation

We investigated the biomass, vertical distribution, and specific root length (SRL) of fine and small roots in a chronosequence of Japanese cedar (Cryptomeria japonica D. Don) plantations in Nara Prefecture, central Japan. Roots were collected from soil blocks up to 50 cm in depth in five plantations of differing age: 4, 15, 30, 41, and 88 years old. Fine-root biomass reached a maximum (639 g m₋₂) in the 15-year-old stand before canopy closure, decreased in the 30-year-old stand (422 g m₋₂), and thereafter was stable. Except in the 30-year-old stand, fine-root biomass increased in deeper soil layers as stand age increased, and the depth at which the cumulative biomass of fine roots reached 90% exhibited a good allometric relationship with mean stem diameter. Both root-length density (root length per unit soil volume) and SRL decreased with soil depth in all stands, indicating that plants mainly acquire water and nutrients from shallow soils. The highest SRL was observed in the 4-year-old stand, but the relationship between SRL and stand age was unclear in older stands. The SRL in surface soils seemed to decrease with increases in root-length density, suggesting that branching of the fine-root system during development is related to density-dependent processes rather than age.     

Estimation of the biomass of fine roots and mycorrhizal fungi: a case study in a Japanese red pine (Pinus densiflora) stand

As part of a study on soil carbon flow in forest ecosystems, the biomass of fine roots (2.0mm in diameter) and root-associated fungi, including ectomycorrhizal fungi, were estimated in the summer season in 1998 at a Pinus densiflora (Japanese red pine) stand in western Japan. Fine roots of pine were classified into three categories: class I roots (0.5–2.0mm in diameter), long class II roots (long roots with diameter 0.5mm; II_L), and short class II roots (short roots with diameter 0.5mm; II_S). Total biomass of fine roots (I + II_L + II_S) at this stand was estimated to be 91.0gm^–2, about 23% of which was class II roots (II_L + II_S). Ergosterol, which is a component of fungal membranes, was analyzed to estimate the biomass of root-associated fungi in roots. In the upper soil layers (from the surface to 13.4cm in depth), ergosterol contents in the class I, II_L and II_S roots were in the ranges 43.1–82.2, 126.1–196.3 and 271.2–321.0µgg^–1 root DW, respectively. The ergosterol content was converted to fungal biomass using the median (minimum–maximum) value of ergosterol concentration reported for ectomycorrhizal fungi. Root-associated fungal biomass in this stand was estimated to be 2.0 (0.5–9.6) gm^–2. The data suggest the biomass of ectomycorrhizal fungi in the P. densiflora stand is small compared with that in other forest ecosystems.     

Effects of plant roots on soil preferential pathways and soil matrix in forest ecosystems

To characterize effects of plant roots on preferential flow(PF),we measured root length density(RLD)and root biomass(RB) in Jiufeng National Forest Park,Beijing,China.Comparisons were made for RLD and RB between soil preferential pathways and soil matrices.RLD and RB declined with the increasing soil depth(0–10,10–20,20–30,30–40,40–50,50–60 cm) in all experimental plots.RLD was greater in soil preferential pathways than in the surrounding soil matrix and was 69.5,75.0 and72.2 % for plant roots of diameter(d) \1,1 \ d \ 3 and3 \ d \ 5 mm,respectively.Fine root systems had the most pivotal influence on soil preferential flow in this forest ecosystem.In all experimental plots,RB content was the sum of RB from soil preferential pathways and the soil matrix in each soil depth.With respect to 6 soil depth gradient(0–10,10–20,20–30,30–40,40–50,50–60 cm) in each plot,the number of soil depth gradient that RB content was greater in soil preferential pathways than in the soil matrix was characterized,and the proportion was68.2 % in all plots.     

Dynamics of nitrogen and phosphorus concentrations of fine roots in a mixed forest ofCunninghamia lanceolata andTsoongiodendron odorum

From September 1999 to July 2000, N and P concentrations of fine roots were measured with the method of sequential soil core at bimonthly intervals in a mixed forest of Tsoong's tree (Tsoongiodendron odorum Chun) and Chinese fir (Cunninghamia lanceolata (Lamb.) Hook.) in Sanming, Fujian. The results showed that N, P concentration of Chinese fir and Tsoong's tree in fine roots were negatively related to root diameter size. The concentrations of N and P in living roots and dead roots were compared. The order of N concentration in fine roots in different samples was Tsoong's tree>undergrowth>Chinese fir, while that of P was undergrowth>Tsoong's tree>Chinese fir. For Chinese fir, the seasonal change of N, P concentrations in fine roots with various diameter classes showed a single-apex curve with a maximum in September. For Tsoong's tree, maximized concentration of N in fine roots appeared in July or September and maximized P concentration in May. Foundation item: The project was supported by The Foundation of Post-doctoral Research of China (1999, No 10), the Foundation for University Key Teacher by the Ministry of Education, and the Research Programs on Basic Theory of Fujian Province (2000F004). Biography: YANG Yu-sheng (1964-), male, professor in Fujian Agricultural and Forestry University, Nanping 353001, P.R. China. Responsible editor: Zhu Hong     

Root parameters of forest trees as sensitive indicators of acidifying pollutants: a review of research of Japanese forest trees

The root parameters of forest trees can be indicators of a changing environment. We summarize the results of root studies with regard to the effects of acidifying pollutants, especially soil acidification and aluminum toxicity, on various root parameters of Japanese forest trees under experimentally controlled conditions. All root parameters such as biomass, morphology, nutritional status, and physiology can be regarded as indicators, because, under laboratory conditions, root responses occur prior to the responses in the aboveground parts. However, considering the conditions of forest sites, the nutritional status and physiological changes are better indicators of soil acidification and Al stress than the biomass and morphological response. The currently available data suggest that the most important indicator is the Ca/Al molar ratio in roots of Japanese tree species. In order to predict and detect the initial effects of soil acidification, we postulate that the specific root response to the Ca/Al molar ratio of tree roots should be considered as a parameter for use in long-term forest monitoring sites.     

Life cycles of individual roots in fine root system of Chamaecyparis obtusa Sieb. et Zucc.

Recent studies have remarked on differences in the life cycles of individual fine roots. However, the dynamics of individual roots with different life cycles, such as ephemeral and perennial, during root system development are still unknown. We examined individual roots during fine root system development in a mature stand of Chamaecyparis obtusa Sieb. et Zucc. (Cupressaceae) using the sequential ingrowth core method and an anatomical method. The visual classification, i.e., orange, red, brown, intact dead, and fragmented dead, of fine roots corresponded well with the anatomical classification. Orange and red roots contained passage cells, and brown roots contained cork cambium. The proportions of protoxylem groups differed among visual classes. Brown secondary roots were mainly triarch (43%) and tetrarch (40%) and rarely diarch (12%), whereas fragmented dead roots, which constituted more than 95% of the dead roots, were mainly diarch (67%). These results imply that triarch and tetrarch roots tend to form secondary roots, whereas diarch roots tend to become dead roots without secondary growth. Using the numbers of root tips and clusters, root system development could be classified into three stages: colonization, branching within the root system, and maintenance. During the colonization stage, mainly triarch and tetrarch roots, which tend to be secondary growth, invaded ingrowth cores. During the branching stage, primarily diarch roots, which tend to be ephemeral, emerged. Fine root system development involved the recruitment of different individual roots during the life cycle depending on the growth stage.     

Effects of site management on growth,biomass partitioning and light use efficiency in a young stand of Eucalyptus grandis in South Africa

The effects of intensive site management treatments at establishment on the production ecology of a stand of Eucalyptus grandis were evaluated in South Africa. Treatments mimicked common operational practices in the region, and included slash removal, slash conservation, slash burning, topsoil disturbance through mechanised harvesting and fertilisation. We calculated the carbon distribution in the standing biomass from allometric relationships. Fine root turnover and litterfall measurements were determined using sequential coring techniques and litter traps, respectively, and this data was used to construct a full model of biomass allocation among stand components. Differences in nutrient availability to young trees, brought about by the most extreme site management treatments, produced several small but significant changes in the elements of the system's production ecology: Absorbed photosynthetically active radiation (APAR) increased from 210 to 247 Mmol photons ha⁻¹ over the 3-year monitoring period, apparent canopy quantum efficiency (α; defined as gross primary production per unit of APAR) from 0.026 to 0.029 mol C (mol photon)⁻¹, and the fraction of carbon allocated to stem wood from 32.7% to 35.6% of net primary production. The magnitudes of these individual responses collectively described the increase in net primary productivity and the Type 1 timber volume response obtained. The biggest changes occurred in APAR, in contrast to published studies from higher rainfall environments where differences in nutrient availability caused greater changes in α than in APAR.     

Above- and below-ground biomass dynamics in a sole cropping and an alley cropping system withGliricidia sepium in the semi-deciduous rainforest zone of West Africa

A considerable amount of data is available about above-ground biomass production and turnover in tropical agroforestry systems, but quantitative information concerning root turnover is lacking. Above- and below-ground biomass dynamics were studied during one year in an alley cropping system withGliricidia sepium and a sole cropping system, on aPlinthic Lixisol in the semi-deciduous rainforest zone of the Côte d'Ivoire. Field crops were maize and groundnut. Live root mass was higher in agroforestry than in sole cropping during most of the study period. This was partly due to increased crop and weed root development and partly to the presence of the hedgerow roots. Fine root production was higher in the alleys and lower under the hedgerows compared to the sole cropping plots. Considering the whole plot area, root production in agroforestry and sole cropping systems was approximatly similar with 1000–1100 kg ha^–1 (dry matter with 45% C) in 0–50 cm depth; about 55% of this root production occured in the top 10 cm. Potential sources of error of the calculation method are discussed on the basis of the compartment flow model. Above-ground biomass production was 11.1 Mg ha^–1 in sole cropping and 13.6 Mg ha^–1 in alley cropping, of which 4.3 Mg ha^–1 were hedgerow prunings. The input of hedgerow root biomass into the soil was limited by the low root mass ofGliricidia as compared to other tree species, and by the decrease of live root mass of hedgerows and associated perennial weeds during the cropping season, presumably as a result of frequent shoot pruning.     

水曲柳落叶松混交林中细根空间分布

采用根钻取样方法对年生水曲柳落叶松混交林中细根空间分布状况进行了研究。结果表明,水曲柳落叶松地下生物量的空间分配差异显著。在林分水平上,水曲柳的根生物量密度高于落叶松(分别为4442.3和2234.9g/m3)。两树种在相邻区域中分配的细根生物量较高,表明种间根系竞争较弱。落叶松行间的水曲柳细根生物量密度和根长密度均高于水曲柳行间的落叶松细根,表明水曲柳地下部分具有较强能力。根系的空间分布有利于混交林中水曲柳的生长。图1表4参19。     

The future of forest research in a changing world

New challenges are facing the managers of the world’s forests, with stakeholders demanding a broader range of goods and services. Balancing the demand for forest products and the responsibility for forest protection is not an easy task. The earlier narrow perspective of wood-resources sustainability has often been transferred to an equally narrow biological diversity conservation perspective. Segregation, in which most of the fiber is produced in plantations, attempts to solve the conflict between conservation and wood production. Biotechnology offers a strategy to gain more wood on less land and with less harm to the environment. The traditional forest research community has not always been able to react properly and promptly to the needs of the users of research results. Consequently, forest research has lost ground to other disciplines. Forest researchers should not consider the newcomers as competitors but instead should try to create more collaboration with those who are interested in solving forest-related problems but do not belong to the old forestry family. There is also a clear need to improve the interface between the research and user communities. Very often the problem is not so much a need to obtain more data than in discovering how to find the most appropriate existing information. In general, forest research should be more cost-efficient. Adopting a more business-like environment should not, however, lead to an excessively result-oriented, short-term way of thinking, to the extent that basic research, and also quality of research, would be in danger.     

Estimated stocks of organic carbon in mangrove roots and sediments in Hinchinbrook Channel,Australia

Aboveground and belowground root biomasses (Babove and Broot) were measured for young, isolated Rhizophorastylosa on Iriomote Island, Japan. The relationship between these two parameters was significant and given as the equation, Broot(g dry weight) = 0.394 × Babove(g dry weight) – 485 (r = 0.986). Multiple regression analyses also revealed good correlation between diameter and biomass of prop roots (Dprop and Bprop) and between prop root and root biomasses. Consequently, root biomass could be estimated from the measurements of diameter and biomass of prop roots using the multiple regression equation, Broot(g dry weight) = 80.0 ×Dprop(cm) + 0.86 ×Bprop (g dry weight) – 251. The relationship between DBH (diameter at breast height) and prop root biomass was also adequately described using an allometric equation.In Hinchinbrook Channel, Australia, redox potential (measured as Eh) and organic carbon stocks in the top 5cm of mangrove sediments were measured along a 600m transect from the frequently inundated, Rhizophora dominated zone on the creek edge, towards higher grounds, where Ceriops spp. became increasingly dominant. Eh values were about –60mV near the creek edge and increased to 260mV on higher grounds. Organic carbon stocks showed an opposite trend to Eh, with the values decreasing from about 360tCha–1 to 160tCha–1. At 18 sites, representing six different habitats, organic carbon stocks were also measured along with the DBH of mangrove trees. DBH was converted into aboveground biomass and then into root biomass using the equations obtained in the study on Iriomote Island. The average organic carbon stocks in the top 50 cm of sediments, aboveground biomass and root biomass were 296tCha–1, 123 tCha–1 and 52 tCha–1, respectively, and accounted for 64%, 25% and 11% of the total organic carbon stock.     

Relationship of water to adventitious rooting in stem cuttings of Populus species

Populus species, characterized by fast growth and easy vegetative propagation, are widely used in agroforestry practices. The substantial water requirement of poplars make them interesting subjects for water balance studies. No information exists on soil moisture requirements for initial root and shoot growth of Populus cuttings. This study on leafless hardwood cuttings of Populus x euramericana (Dode) Guinier cv. Robusta examined the dynamics of water use during propagation, as influenced by two initial soil water potentials (–0.006 and –0.06 MPa). Differences in the initial water potential of the cuttings was achieved by three pretreatments i.e., fresh, soaked and dried. Initial _shoot was –1.45, –0.10 and –2.10 MPa in fresh, soaked and dried cuttings, respectively. Soil moisture had a major effect on rooting. Water-stressed cuttings took a longer time to root and had fewer roots. Pre-soaking of cuttings stimulated rooting, particularly under the drier soil moisture conditions. Initially the water potential of cuttings decreased with time and with the formation of roots it stabilized in all the pretreatments. The reduction in water potential of cuttings after planting was related to an increase in resistance to water flow in the xylem.This revised version was published online in November 2005 with corrections to the Cover Date.     

Carbon unloading in roots in relation to root senescence in Cercis chinensis seedlings under drought stress

When Cercis chinensis seedlings suffered from drought treatment,net photosynthetic rates had been significantly reduced at the end of the drought treatment. Compared with the control,the activities of acid invertases in roots had increased 5 and 11 days after drought treatment. Seventeen days after drought treatment,the activities of acid invertases in roots were significantly decreased,while activities of alkaline invertases in roots had also been significantly reduced. As the moisture in culture media dec...