Competition in tree row agroforestry systems. 1. Distribution and dynamics of fine root length and biomass

Complementarity in the distribution of tree and crop root systems is important to minimise competition for resources whilst maximising resource use in agroforestry systems. A field study was conducted on a kaolinitic Oxisol in the sub-humid highlands of western Kenya to compare the distribution and dynamics of root length and biomass of a 3-year-old Grevillea robusta A. Cunn. ex R. Br. (grevillea) tree row and a 3-year-old Senna spectabilis DC. (senna) hedgerow grown with Zea mays L. (maize). Tree roots were sampled to a 300 cm depth and 525 cm distance from the tree rows, both before and after maize cropping. Maize roots were sampled at two distances from the tree rows (75–150 cm and 450–525 cm) to a maximum depth of 180 cm, at three developmental stages. The mean root length density (L_rv) of the trees in the upper 15 cm was 0.55 cm cm⁻³ for grevillea and 1.44 cm cm⁻³ for senna, at the start of the cropping season. The L_rv of senna decreased at every depth during the cropping season, whereas the L_rv of grevillea only decreased in the crop rooting zone. The fine root length of the trees decreased by about 35% for grevillea and 65% for senna, because of maize competition, manual weeding, seasonal senescence or pruning regime (senna). At anthesis, the L_rv of maize in the upper 15 cm was between 0.8 and 1.5 cm cm⁻³. Maize root length decreased with greater proximity to the tree rows, potentially reducing its ability to compete for soil resources. However, the specific root length (m g⁻¹) of maize was about twice that of the trees, so may have had a competitive uptake advantage even when tree root length was greater. Differences in maize fine root length and biomass suggest that competition for soil resources and hence fine root length may have been more important for maize grown with senna than grevillea. This revised version was published online in June 2006 with corrections to the Cover Date.     

Annual and seasonal changes in fine root biomass of a Quercus ilex L. forest

The biomass, production and mortality of fine roots (roots with diameter <2.5 mm) were studied in a typical Mediterranean holm oak (Quercus ilex L.) forest in NE Spain using the minirhizotron methodology. A total of 1212 roots were monitored between June of 1994 and March of 1997. Mean annual fine root biomass in the holm oak forest of Prades was 71±8 g m^–2 yr^–1. Mean annual production for the period analysed was 260+11 g m^–2 yr^–1. Mortality was similar to production, with a mean value of 253±3 g m^–2 yr^–1. Seasonal fine root biomass presented a cyclic behaviour, with higher values in autumn and winter and lower in spring and summer. Production was highest in winter, and mortality in spring. In summer, production and mortality values were the lowest for the year. Production values in autumn and spring were very similar. The vertical distribution of fine root biomass decreased with increasing depth except for the top 10–20 cm, where values were lower than immediately below. Production and mortality values were similar between 10 and 50 cm depth. In the 0–10 cm and the 50–60 cm depth intervals, both production and mortality were lower.     

Root production and root turnover in two dominant species of wet heathlands

Summary Root biomass production, root length production and root turnover of Erica tetralix and Molinia caerulea were estimated by sequential core sampling and by observations in permanent minirhizotrons in the field. Root biomass production, estimated by core sampling, was 370 (Erica) and 1080 (Molinia) g m^-2 yr^-1. This was for both species equal to aboveground production. Assuming steady-state conditions for the root system, root biomass turnover rates (yr^-1), estimated by core sampling, were 1.72 (Erica) and 1.27 (Molinia). Root length production of both species, estimated by minirhizotron observations, varied significantly with observation depth. Root length turnover rate (yr^-1) of both species did not vary significantly with observation depth and averaged 0.92 in Erica and 2.28 in Molinia. Reasons are given for the discrepancy between the results of the two types of turnover measurements. The data suggest that the replacement of Erica by Molinia in a wet heathland, which occurs when nutrient availability increases, leads to an increased flow of carbon and nutrients into the soil-system. Therefore, there may be a positive feedback between dominance of Molinia and nutrient availability.     

毛竹细根分布特征研究

为了解毛竹(Phyllostachys edulis)细根的分布规律,对不同水平距离和土层深度0~1 mm和1~2 mm细根的生物量、比根长、组织密度和根长密度进行了分析。结果表明,随着毛竹年龄的增加,细根生物量和根长密度先上升后降低,根组织密度先降低后升高,比根长呈降低的趋势。细根生物量和根长密度以距竹秆60 cm处最大,根组织密度以20 cm处最大,比根长在40 cm处最大,但他们在距竹秆不同距离间的差异不显著。细根生物量以10~20 cm土层最大,根组织密度以20~30 cm土层最大,细根生物量、比根长、组织密度和根长密度在不同土层间的差异不显著。与1~2 mm细根相比,0~1 mm细根生物量和根组织密度更小,比根长和根长密度更大。因此,毛竹年龄对细根生长具有显著的影响,1年生毛竹有最大的比根长和较大的根组织密度,具有更强的资源利用率。毛竹细根在一定的土层范围内呈均匀分布状态,可更有效地利用特定区域的水肥资源。     

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]     

Fibrous carrot root responses to irrigation and compaction of sandy and organic soils

Field experiments were performed in Southern Finland on fine sand and organic soil in 1990 and 1991 to study carrot roots. Fall ploughed land was loosened by rotary harrowing to a depth of 20 cm or compacted under moist conditions to a depth of 25–30 cm by three passes of adjacent wheel tracks with a tractor weighing 3 Mg, in April were contiguously applied across the plot before seed bed preparation. Sprinkler irrigation (30 mm) was applied to fine sand when moisture in the 0–15 cm range of soil depth was 50% of plant-available water capacity. For root sampling, polyvinyl chloride (PVC) cylinders (30 × 60 cm) were installed in the rows of experimental plots after sowing, and removed at harvest. Six carrot plants were grown in each of in these soil colums in situ in the field.Fine root length and width were quantified by image analysis. Root length density (RLD) per plant was 0.2–1.0 cm cm^-3 in the 0–30 cm range. The fibrous root system of one carrot had total root lengths of 130–150 m in loose fine sand and 180–200 m in compacted fine sand. More roots were observed in irrigated than non-irrigated soils. In the 0–50 cm range of organic soil, 230–250 m of root length were removed from loosened organic soils and 240–300 m from compacted soils. Specific root surface area (surface area divided by dry root weight) of a carrot fibrous root system averaged 1500–2000 cm² g^-1. Root length to weight ratios of 250–350 m g^-1 effectively compare with the ratios of other species.Fibrous root growth was stimulated by soil compaction or irrigation to a depth of 30 cm, in both the fine sand and organic soils, suggesting better soil water supply in compacted than in loosened soils. Soil compaction increased root diameters more in fine sand than it did in organic soil. Most of the root length in loosened soils (fine sand 90%, organic soil 80%) and compacted soils (fine sand 80%, organic soil 75%) was composed of roots with diameters of approximately 0.15 mm. With respect to dry weight, length, surface area and volume of the fibrous root system, all the measurements gave significant resposes to irrigation and soil compaction. Total root volumes in the 0–50 cm of soil were 4.3 cm³ and 9.8 cm³ in loosened fine sand and organic soils, respectively, and 6.7 cm³ and 13.4 cm³ in compacted sand and organic soils, respectively. In fine sand, irrigation increased the volume from 4.8 to 6.3 cm³.     

间伐对黄龙山油松中龄林细根空间分布和形态特征的影响

为探究油松细根生长与抚育间伐的关系,以黄龙山林区4种不同间伐强度(对照,轻度,中度,强度)下的油松人工中龄林为研究对象,采用根钻法,分3层(0—20,20—40,40—60cm)获取细根样品,研究了间伐强度对油松细根生物量和形态特征的影响。结果表明:油松细根生物量主要分布在0—20 cm土层,不同间伐强度下细根生物量差异显著(P0.05),随间伐强度的增大,细根生物量先升高后降低,强度间伐下0—20 cm土层细根生物量显著降低(P0.05),20—40 cm土层和40—60 cm土层细根生物量所占比例随间伐强度的增大而增大。细根根长密度和根表面积密度在不同间伐强度和不同土层间均差异显著(P0.05),且变化规律与生物量基本一致。细根比根长和比表面积随间伐强度的增加而增大,且强度间伐与其他强度呈显著性差异(P0.05)。轻度和中度间伐对小径级细根(0—1.0 mm)有显著影响,对较大径级细根(1.0—2.0 mm)的影响则不显著(P0.05),强度间伐对0—2.0mm的细根均有显著影响(P0.05)。中度间伐(保留郁闭度0.7)条件下,油松林地细根总生物量达到最大1022.43 g/m2,此条件下细根的根长密度和根表面积密度也达到最大,能充分利用林地的立地资源,最有利于保留木的生长。     

青海省森林细根生物量及其影响因子

细根是植物吸收水分和养分的主要器官。全球变暖背景下,研究森林细根生物量及其环境因子的变化对生态系统碳平衡、碳收支及其贡献率具有重要意义。采用土钻法和室内分析法对青海省森林6个海拔梯度上5种林分类型的细根生物量和土壤理化性质进行测定,并分析了与环境因子之间的相互关系。结果表明:(1)青海省森林0—40 cm土层总细根生物量平均为8.50 t/hm~2,随着海拔梯度的增加先降低后升高,不同海拔梯度细根生物量差异显著(P0.05),最大值出现在2100—2400 m处。(2)5种林分0—40 cm土层总细根生物量为:白桦白杨云杉圆柏山杨,不同林分间细根生物量差异不显著。(3)细根垂直分布随土层深度增加而减少,且70%的细根集中在表层(0—20 cm)。(4)土壤容重深层(20—40 cm)显著大于表层(P0.05),并随海拔梯度逐步增加,且林分间差异较大。(5)全碳(Total carbon, TC)、全氮(Total nitrogen, TN)、全磷(Total phosphorus, TP)含量表层显著高于深层。TC、TN随海拔升高先增后降低,TP则随海拔逐步降低。不同林分间土壤养分差异较明显。(6)结构方程模型分析得到海拔、土层、容重直接影响细根生物量,细根生物量直接影响土壤养分。林分类型通过土壤容重间接影响细根生物量。因此,林分和海拔通过影响土壤微环境而影响到细根生物量及其空间分布格局。     

Root biomass changes after drainage and fertilization of a low-shrub pine bog

The stump and root systems of Scots pine (Pinus sylvestris) and field-layer vegetation were sampled before (1984) and three growing seasons after drainage and fertilization (1987) of a low-shrub pine bog. Average below-ground biomass of the field layer was 548 gDW m^–2 in 1984, with no significant treatment effects during experimentation. The stump-plus-root biomass of the pine stands was 1464 gDW m^–2 in the virgin state, and had increased to 1854 gDW m^–2 three years after the NPK-fertilizer treatment. The distribution over fractions also changed with this treatment. The fraction of fine roots ( < 1 mm) in stump-root biomass increased from 4% (56 gDW m^–2) to 11% (196 gDW m^–2), while the other compartments changed less. Total pine root length was 729 mm^–2 in 1984. Root length increased by 94% to 1380 mm^–2 on NPK-fertilized plots. Most of the fine pine roots were in the surface layer (0–10 cm), 79% in 1984 and 88% in 1987, and few pine roots were deeper than 20 cm. Maximum root length of fine pine roots ( < 1 mm) was estimated to be 2710 mm^–2 at about 800 gDW m^–2 (NPK treatment), and the corresponding maximum for small pine roots (=1–10 mm) was 227 mm^–2 at 809 gDW m^–2. Drainage stimulated net growth of fine roots, but this treatment also caused higher mortality rates of small roots. The fine roots responded to fertilization with higher net growth rate, and secondary growth of the large roots ( > 10 mm) was improved. The observed changes in root biomass and structure are explained as strategic adaptations to altered hydrological and nutritional circumstances in the root zone after drainage and fertilization.     

Fine root demography in alfalfa (Medicago sativa L.)

In perennial forages like alfalfa (Medicago sativa L.), repeated herbage removal may alter root production and mortality which, in turn, could affect deposition of fixed N in soil. Our objective was to determine the extent and patterns of fine-diameter root production and loss during the year of alfalfa stand establishment. The experiment was conducted on a loamy sand soil (Udorthentic Haploboroll) in Minnesota, USA, using horizontally installed minirhizotrons placed directly under the seeded rows at 10, 20, and 40 cm depths in four replicate blocks. We seeded four alfalfa germplasms that differed in N₂ fixation capacity and root system architecture: Agate alfalfa, a winter hardy commercially-available cultivar; Ineffective Agate, which is a non-N₂-fixing near isoline of Agate; a new germplasm that has few fibrous roots and strong tap-rooted traits; and a new germplasm that has many fibrous roots and a strongly branched root system architecture. Video images collected biweekly throughout the initial growing season were processed using C-MAP-ROOTS software.More than one-half of all fine roots in the upper 20 cm were produced during the first 7 weeks of growth. Root production was similar among germplasms, except that the highly fibrous, branch-rooted germplasm produced 29% more fine roots at 20 cm than other germplasms. In all germplasms, about 7% of the fine roots at each depth developed into secondarily thickened roots. By the end of the first growing season, greatest fine root mortality had occurred in the uppermost depth (48%), and least occurred at 40 cm (36%). Survival of contemporaneous root cohorts was not related to soil depth in a simple fashion, although all survivorship curves could be described using only five rates of exponential decline. There was a significant reduction in fine root mortality before the first herbage harvest, followed by a pronounced loss (average 22%) of fine roots at the 10- and 20-cm depths in the 2-week period following herbage removal. Median life spans of these early-season cohorts ranged from 58 to 131 days, based on fitted exponential equations. At all depths, fine roots produced in the 4 weeks before harvest (early- to mid-August) tended to have shorter median life spans than early-season cohorts. Similar patterns of fine root mortality did not occur at the second harvest. Germplasms differed in the pattern, but not the ultimate extent, of fine root mortality. Fine root turnover during the first year of alfalfa establishment in this experiment released an estimated 830 kg C ha^–1 and 60 kg N ha^–1, with no differences due to N₂ fixation capacity or root system architecture.     

The effect of increased nutrient availability on vegetation dynamics in wet heathlands

A three year fertilization experiment was conducted in which nitrogen (N series: 20 g N m^–2 yr^–1), phosphorus (P series: 4 g P m^–2 yr^–1) and potassium (K series: 20 g K m^–2 yr^–1) were added to a mixed vegetation of Erica tetralix and Molinia caerulea. At the end of each growing season the percentage cover of each species was determined. At the end of the experiment percentage cover of each species was found to be positively correlated with the harvested biomass. In the unfertilized control series the cover of Erica and Molinia did not change significantly during the experiment. In all fertilized series however, especially in the P series, cover of Erica decreased significantly. The cover of Molinia increased significantly in the P series only.In the fertilized series the biomass of Erica and total biomass per plot did not change significantly compared with the control series. In the P series the biomass of Molinia increased significantly.It is suggested that with increasing phosphorus or nitrogen availability Molinia outcompetes Erica because the former invests more biomass in leaves which in turn permits more carbon to be allocated to the root system, which thereupon leads to a higher nutrient uptake.     

Morphological synergism in root hair length,density, initiation and geometry for phosphorus acquisition in Arabidopsis thaliana: A modeling approach

Low phosphorus availability regulates root hair growth in Arabidopsis by (1) increasing root hair length, (2) increasing root hair density, (3) decreasing the distance between the root tip and the point at which root hairs begin to emerge, and (4) increasing the number of epidermal cell files that bear hairs (trichoblasts). The coordinated regulation of these traits by phosphorus availability prompted us to speculate that they are synergistic, that is, that they have greater adaptive value in combination than they do in isolation. In this study, we explored this concept using a geometric model to evaluate the effect of varying root hair length (short, medium, and long), density (0, 24, 48, 72, 96, and 120 root hairs per mm of root length), tip to first root hair distance (0.5, 1, 2, and 4 mm), and number of trichoblast files (8 vs. 12) on phosphorus acquisition efficiency (PAE) in Arabidopsis. SimRoot, a dynamic three-dimensional geometric model of root growth and architecture, was used to simulate the growth of Arabidopsis roots with contrasting root hair parameters at three values of phosphorus diffusion coefficient (D _e=1×10^–7, 1×10^–8, and 1×10^–9 cm² s^–1) over time (20, 40, and 60 h). Depzone, a program that dynamically models nutrient diffusion to roots, was employed to estimate PAE and competition among root hairs. As D _e decreased from 1×10^–7 to 1×10^–9 cm² s^–1, roots with longer root hairs and higher root hair densities had greater PAE than those with shorter and less dense root hairs. At D _e=1×10^–9 cm² s^–1, the PAE of root hairs at any given density was in the order of long hairs > medium length hairs > short hairs, and the maximum PAE occurred at density = 96 hairs mm^–1 for both long and medium length hairs. This was due to greater competition among root hairs when they were short and dense. Competition over time decreased differences in PAE due to density, but the effect of length was maintained, as there was less competition among long hairs than short hairs. At high D _e(1×10^–7 cm² s^–1), competition among root hairs was greatest among long hairs and lowest among short hairs, and competition increased with increasing root hair densities. This led to a decrease in PAE as root hair length and density increased. PAE was also affected by the tip to first root hair distance. At low D _e values, decreasing tip to first root hair distance increased PAE of long hairs more than that of short hairs, whereas at high D _e values, decreasing tip to first root hair distance increased PAE of root hairs at low density but decreased PAE of long hairs at very high density. Our models confirmed the benefits of increasing root hair density by increasing the number of trichoblast files rather than decreasing the trichoblast length. The combined effects of all four root hair traits on phosphorus acquisition was 371% greater than their additive effects, demonstrating substantial morphological synergy. In conclusion, our data support the hypothesis that the responses of root hairs to low phosphorus availability are synergistic, which may account for their coordinated regulation.     

Fine root architecture, morphology, and biomass of different branch orders of two Chinese temperate tree species

We have limited understanding of architecture and morphology of fine root systems in large woody trees. This study investigated architecture, morphology, and biomass of different fine root branch orders of two temperate tree species from Northeastern China—Larix gmelinii Rupr and Fraxinus mandshurica Rupr —by sampling up to five fine root branch orders three times during the 2003 growing season from two soil depths (i.e., 0–10 and.10–20 cm). Branching ratio (R _b) differed with the level of branching: R _b values from the fifth to the second order of branching were approximately three in both species, but markedly higher for the first two orders of branching, reaching a value of 10.4 for L. gmelinii and 18.6 for F. mandshurica. Fine root diameter, length, SRL and root length density not only had systematic changes with root order, but also varied significantly with season and soil depth. Total biomass per order did not change systematically with branch order. Compared to the second, third and/or fourth order, the first order roots exhibited higher biomass throughout the growing season and soil depths, a pattern related to consistently higher R _b values for the first two orders of branching than the other levels of branching. Moreover, the differences in architecture and morphology across order, season, and soil depth between the two species were consistent with the morphological disparity between gymnosperms and angiosperms reported previously. The results of this study suggest that root architecture and morphology, especially those of the first order roots, should be important for understanding the complexity and multi-functionality of tree fine roots with respect to root nutrient and water uptake, and fine root dynamics in forest ecosystems.     

Biomass and distribution of fine and coarse roots from blue oak (Quercus douglasii) trees in the northern Sierra Nevada foothills of California

This research adds to the limited data on coarse and fine root biomass for blue oak (Quercus douglasii Hook and Arn.), a California deciduous oak species found extensively throughout the interior foothills surrounding the Central Valley. Root systems of six blue oak trees were analyzed using three methods — backhoe excavation, quantitative pits, and soil cores. Coarse root biomass ranged from 7 to 177 kg per tree. Rooting depth for the main root system ranged from 0.5 to 1.5 m, with an average of 70% of excavated root biomass located above 0.5 m. Of the total biomass in excavated central root systems, primary roots (including burls) accounted for 56% and large lateral roots (> 20 mm diameter) accounted for 36%. Data from cores indicated that most biomass outside of the root crown was located in fine roots and that fine root biomass decreased with depth. At surface depths (0–20 cm), small-fine (< 0.5 mm diameter) roots accounted for 71%, large-fine (0.5–2.0 mm) for 25%, and coarse (> 2 mm) for 4% of total root biomass collected with cores. Mean fine root biomass density in the top 50 cm was 0.43 kg m⁻³. Fine root biomass did not change with increasing distance from the trees (up to approximately 5 m). Thus, fine roots were not concentrated under the tree canopies. Our results emphasize the importance of the smallest size class of roots (<0.5 mm), which had both higher N concentration and, in the area outside the central root system, greater biomass than large fine (0.5–2.0 mm) or coarse (> 2.0 mm) roots. This revised version was published online in June 2006 with corrections to the Cover Date.     

Size and Structure of Fine Root Systems in Old-growth and Secondary Tropical Montane Forests (Costa Rica)

The fine root systems of three tropical montane forests differing in age and history were investigated in the Cordillera Talamanca, Costa Rica. We analyzed abundance, vertical distribution, and morphology of fine roots in an early successional forest (10–15 years old, ESF), a mid‐successional forest (40 years old, MSP), and a nearby undisturbed old‐growth forest (OGF), and related the root data to soil morphological and chemical parameters. The OGF stand contained a 19 cm deep organic layer on the forest floor (i.e., 530 mol C/m²), which was two and five times thicker than that of the MSF (10 cm) and ESF stands (4 cm), respectively. There was a corresponding decrease in fine root biomass in this horizon from 1128 g dry matter/m² in the old‐growth forest to 337 (MSF) and 31 g/m² (ESF) in the secondary forests, although the stands had similar leaf areas. The organic layer was a preferred substrate for fine root growth in the old‐growth forest as indicated by more than four times higher fine root densities (root mass per soil volume) than in the mineral topsoil (0–10 cm); in the two secondary forests, root densities in the organic layer were equal to or lower than in the mineral soil. Specific fine root surface areas and specific root tip abundance (tips per unit root dry mass) were significantly greater in the roots of the ESF than the MSF and OGF stands. Most roots of the ESF trees (8 abundant species) were infected by VA mycorrhizal fungi; ectomycorrhizal species (Quercus copeyemis and Q. costaricensis) were dominant in the MSF and OGF stands. Replacement of tropical montane oak forest by secondary forest in Costa Rica has resulted in (1) a large reduction of tree fine root biomass; (2) a substantial decrease in depth of the organic layer (and thus in preferred rooting space); and (3) a great loss of soil carbon and nutrients. Whether old–growth Quercus forests maintain a very high fine root biomass because their ectomycorrhizal rootlets are less effective in nutrient absorption than those of VA mycorrhizal secondary forests, or if their nutrient demand is much higher than that of secondary forests (despite a similar leaf area and leaf mass production), remains unclear.     

Alley cropping with Senna siamea in South-western Nigeria: II. Dry matter,total N,and urea-derived N dynamics of the Senna and maize roots

Crop and tree roots are crucial in the nutrient recycling hypotheses related to alley cropping systems. At the same time, they are the least understood components of these systems. The biomass, total N content and urea-derived N content of the Senna and maize roots in a Senna-maize alley cropping system were followed for a period of 1.5 years (1 maize-cowpea rotation followed by 1 maize season) to a depth of 90 cm, after the application of ¹⁵N labeled urea. The highest maize root biomass was found in the 0–10 cm layer and this biomass peaked at 38 and 67 days after planting the 1994 maize (DAP) between the maize rows (112 kg ha^–1, on average) and at 38, 67 and 107 DAP under the maize plants (4101 kg ha^–1, on average). Almost no maize roots were found below 60 cm at any sampling date. Senna root biomass decreased with time in all soil layers (from 512 to 68 kg ha^–1 for the 0–10 cm layer between 0 and 480 DAP). Below 10 cm, at least 62% of the total root biomass consisted of Senna roots and this value increased to 87% between 60 and 90 cm. Although these observations support the existence of a Senna root `safety net' between the alleys which could reduce nutrient leaching losses, the depth of such a net may be limited as the root biomass of the Senna trees in the 60–90 cm layer was below 100 kg ha^–1, equivalent to a root length density of only < 0.05 cm cm^–3. The proportion of maize root N derived from the applied urea (%Ndfu) decreased significantly with time (from 21% at 21 DAP to 8% at 107 DAP), while %Ndfu of the maize roots at the second harvest (480 DAP) was only 0.6%. The %Ndfu of the Senna roots never exceeded 4% at any depth or sampling time, but decreased less rapidly compared to the %Ndfu of the maize roots. The higher %Ndfu of the maize roots indicates that maize is more efficient in retrieving urea-derived N. The differences in dynamics of the %Ndfu also indicate that the turnover of N through the maize roots is much faster than the turnover of N through the Senna roots. The recovery of applied urea-N by the maize roots was highest in the top 0–10 cm of soil and never exceeded 0.4% (at 38 DAP) between the rows and 7.1% (at 67 DAP) under the rows. Total urea N recovery by the maize roots increased from 1.8 to 3.2% during the 1994 maize season, while the Senna roots never recovered more than 0.8% of the applied urea-N at any time during the experimental period. These values are low and signify that the roots of both plants will only marginally affect the total recovery of the applied urea-N. Measurement of the dynamics of the biomass and N content of the maize and Senna roots helps to explain the observed recovery of applied urea-N in the aboveground compartments of the alley cropping system.     

杉木叶片、细根功能性状对毛竹扩张及伐除的响应

功能性状能够反映植物对不同环境的适应策略。毛竹扩张与外来植物入侵相似，常引起原有植物生存环境的改变，而原有植物功能性状对毛竹扩张及伐除的响应机制尚不清楚。选取毛竹-杉木混交林和去竹杉木林为研究对象，以杉木纯林为对照，比较分析杉木比叶面积、叶干物质含量、叶组织密度等叶功能性状以及比根长、细根生物量、细根根长密度等细根功能性状的变化以及其间的相关关系。结果表明：(1)与杉木纯林相比，混交林中杉木的叶相对含水量以及叶干物质含量分别减少了5.07%、0.032 g/g,叶组织密度以及比叶面积分别增加了0.005 g/cm³、10.33 cm²/g;而去竹杉木林中，杉木比叶面积、叶相对含水量减少，叶干物质含量和叶组织密度则呈上升趋势。(2)与杉木纯林相比，混交林中杉木细根生物量、细根体积密度以及细根根长密度都不断下降，而杉木细根比根长在0—20 cm土深处显著增加(P<0.05);而去竹杉木林中杉木细根比根长、细根根长密度和细根生物量则显著降低(P<0.05),细根体积密度在20—30 cm土深处有所增加。(3)杉木纯林中杉木细根功能性状间...     

间伐对川西亚高山粗枝云杉人工林细根生物量及碳储量的影响

以川西亚高山50年生粗枝云杉(Picea asperata)人工林为研究对象, 探讨了间伐对粗枝云杉人工林1-5级细根生物量及碳储量的影响。结果表明: 粗枝云杉人工林细根生物量和碳储量随根序等级的增加而显著增加(p < 0.05), 5级根序中1级根生物量及碳储量最小, 5级根生物量及碳储量最大。与对照(间伐0%)相比, 间伐对粗枝云杉人工林林分细根生物量及碳储量有显著影响(p < 0.05); 而对单株细根生物量影响不一, 间伐10%和20%与对照没有显著性差异(p > 0.05)。间伐显著影响生物量在各根序中的分配, 随着间伐强度的增加, 1、2级细根中生物量分配比例增加, 1级细根的生物量增加幅度最大; 3-5级细根的生物量分配比例减小, 5级细根减少幅度最大。其中, 间伐50%显著减少了细根在下层(20-40 cm)土壤中的生物量比例(p < 0.05), 但与间伐20%和30%无显著差异(p > 0.05)。     

不同林龄麻栎林地下部分生物量与碳储量研究

探讨不同林龄麻栎林地下部分根系的生物量与碳储量,为麻栎林的经营管理及碳汇管理等提供科学依据。以江苏省句容市不同林龄(幼龄林、中龄林、近熟林、成熟林)的麻栎林为研究对象,采用全根挖掘法获取麻栎各级根系及灌草层根系,并测定其生物量、碳含量,构建麻栎根系生物量模型,估算麻栎林地下部分根系碳储量及麻栎林群落碳储量。通过11种数学回归模型的比较,构建麻栎各级根系生物量幂回归模型,计算得到幼龄林、中龄林、近熟林、成熟林麻栎根系生物量分别为14.81t/hm~2、41.15t/hm~2、50.36t/hm~2、53.75t/hm~2,各级根系生物量大小顺序是:根桩粗根大根细根;灌木与草本植物根系生物量分别为0.48—1.71t/hm~2、0.13—0.60t/hm~2;不同林龄麻栎林群落根系生物量为15.42—56.06t/hm~2,且随林龄的增大而增大。麻栎根系碳含量大小顺序为:根桩粗根大根细根,且碳含量差异显著;灌木与草本植物根系碳含量分别为41.84%—43.79%、34.03%—38.48%,随林龄变化均无明显变化规律。麻栎林乔木根系碳储量随林龄增大而增大,幼龄林、中龄林、近熟林、成熟林根系碳储量分别为6.01t/hm~2、17.41t/hm~2、21.79t/hm~2、21.99t/hm~2;灌木与草本植物根系碳储量均随林龄增大而增大;幼龄林、中龄林、近熟林、成熟林群落根系碳储量分别为6.26t/hm~2、17.74t/hm~2、22.37t/hm~2、22.94t/hm~2,且乔木层灌木层草本层。麻栎林地下部分根系生物量与碳储量随林龄的增大而增大,幼龄林到近熟林生长过程中生物量与碳储量增加快速,近熟林后生物量与碳素积累缓慢,且与成熟林接近。     

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.