Root deformation in plantations of container-grown Scots pine trees: effects on root growth,tree stability and stem straightness

Root system deformation was studied in 23 Scots pine (Pinus sylvestris L.) stands in central Sweden. The study comprised both plantations created with container-grown plants (Paperpot) and natural stands including young (7–9 year old) and older (19–24 year old) trees. Trees were measured with regards to distribution of roots, root deformation, stability, stem straightness and wood properties in stumps. Root distribution was most uniform for naturally regenerated trees. Older trees generally showed a better root distribution than young trees. The young planted trees displayed a high frequency of severely spiralled root systems, while only a few of the older trees had spiralled root systems. No severe root deformations were observed on naturally regenerated trees. Naturally regenerated trees were more stable than those which had been planted. Differences in bending moment, when trees were pulled to an angle of 10°, were considerable between young planted and naturally regenerated trees, but less pronounced for the older trees. Young planted trees had the highest frequency of severely crooked stem bases, while naturally regenerated trees had the straightest mode of growth. Tensile strength in peripheral wood samples of the stumps was substantially lower for planted than for naturally regenerated trees. Strain values to breakage of wood samples, taken from the root collar and the central- and peripheral part of the stump were lower for planted trees. The conclusions from this study are that root distribution, tree stability and stem straightness of planted Paperpot-grown trees will improve after a certain time and approach the state of naturally regenerated trees. As trees grow older, early established crooked stem bases will be compensated by radial growth and the tree will appear straighter. Inside the stem, however, problems may still remain with abnormal fibre direction and compression wood together with inferior root strength due to fibre disturbances as a result of spiralled roots. This revised version was published online in June 2006 with corrections to the Cover Date.     

Root System Asymmetry of Mediterranean Pines

Three groups of Mediterranean pines were examined to describe the development of root symmetry on sites characterized by shallow soils and low water availability. Sampling included: (1) 3-year-old planted seedlings of Pinus halepensis Mill. taken from Sithonia Halkidiki, northern Greece, (2) 5-year-old natural regenerated seedlings of Pinus brutia Ten. taken from Kedrinos Lofos, Thessaloniki and (3) 65-year-old trees of Pinus brutia taken from Kedrinos Lofos, Thessaloniki. Root system symmetry was examined by measuring the number, the diameter, the cross-sectional area (CSA), the root area index (RAI) and the length of the lateral roots of each root system, and by analyzing their distribution around the stem. Above-ground plant symmetry was also estimated. The findings of the study indicated that there was an asymmetric root system in all three groups that is characterized by the concentration of the main laterals along the contour lines instead of uphill or downhill; however, the asymmetry was much higher in the young plants. This asymmetry was not correlated with the above-ground plant growth form, which was found to be symmetric. The asymmetric development of root can be attributed to the shallow soil and the high mechanical resistance of the underground bedrock that stopped the taproot growth, restricted the root penetration in the deeper layers and obliged the roots to elongate towards the surface soil layers, where there is more available water.     

Effects of cropping, nutrition and water supply on accumulation and distribution of biomass and nutrients for apple trees on'M9'root systems

The effects of cropping, nutrition and water supply on accumulation and distribution of biomass and nutrients for apple trees on'M9'root systems were examined using trees growing in lysimeters. For 3-year-old 'Golden Delicious' trees, cropping significantly increased total biomass, although leaf and root biomass were reduced. For 3-year-old'Golden Delicious', 'Coxs Orange' and 'Gloster' trees, cropping only in the third year reduced root biomass by 39 to 45%. Cropping did not affect water use by trees of any cultivar early in the season, but increased water use later in the season for 'Gloster' trees. For trees with fruit, total non-structural carbohydrate (TNC) contents at the end of the season were highest in fruit (up to 1 400 g plant ⁻¹). For trees without fruit, TNC contents were generally highest in the roots (up to 110 g planr ⁻¹). Supplying trees with nutrient solution diluted to 10% of the standard composition increased the root biomass for 'Golden Delicious' trees only, but decreased biomass of above-ground tissues for all cultivars. Reducing the nutrient supply decreased water and nutrient uptake, and reduced nutrient contents within vegetative parts of the tree more than within roots. For 'Golden Delicious' trees, restricting the water supply to 50% or 25% of that consumed by control trees significantly reduced above-ground biomass, but root biomass was not significantly affected. The N, P and K. contents for the trees were also reduced by water stress, due to reduced contents within above-ground organs. Water stress reduced the TNC and starch contents of all tissues, except the roots. These results are discussed in relation to the efficiency of root systems for water and nutrient uptake.     

CASIROZ: Root parameters and types of ectomycorrhiza of young beech plants exposed to different ozone and light regimes

Tropospheric ozone (O(3)) triggers physiological changes in leaves that affect carbon source strength leading to decreased carbon allocation below-ground, thus affecting roots and root symbionts. The effects of O(3) depend on the maturity-related physiological state of the plant, therefore adult and young forest trees might react differently. To test the applicability of young beech plants for studying the effects of O(3) on forest trees and forest stands, beech seedlings were planted in containers and exposed for two years in the Kranzberg forest FACOS experiment (Free-Air Canopy O(3) Exposure System, http://www.casiroz.de ) to enhanced ozone concentration regime (ambient [control] and double ambient concentration, not exceeding 150 ppb) under different light conditions (sun and shade). After two growing seasons the biomass of the above- and below-ground parts, beech roots (using WinRhizo programme), anatomical and molecular (ITS-RFLP and sequencing) identification of ectomycorrhizal types and nutrient concentrations were assessed. The mycorrhization of beech seedlings was very low ( CA. 5 % in shade, 10 % in sun-grown plants), no trends were observed in mycorrhization (%) due to ozone treatment. The number of Cenococcum geophilum type of ectomycorrhiza, as an indicator of stress in the forest stands, was not significantly different under different ozone treatments. It was predominantly occurring in sun-exposed plants, while its majority share was replaced by Genea hispidula in shade-grown plants. Different light regimes significantly influenced all parameters except shoot/root ratio and number of ectomycorrhizal types. In the ozone fumigated plants the number of types, number of root tips per length of 1 to 2 mm root diameter, root length density per volume of soil and concentration of Mg were significantly lower than in control plants. Trends to a decrease were found in root, shoot, leaf, and total dry weights, total number of root tips, number of vital mycorrhizal root tips, fine root (mass) density, root tip density per surface, root area index, concentration of Zn, and Ca/Al ratio. Due to the general reduction in root growth indices and nutrient cycling in ozone-fumigated plants, alterations in soil carbon pools could be predicted.     

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.     

Root system architecture of Quercus pubescens trees growing on different sloping conditions

BACKGROUND AND AIMS: Plant roots' growth direction has important implications for plant development and survival; moreover it plays an effective and vital role in stabilizing weathered soil on a steep slope. The aim of this work was to assess the influence of slope on the architecture of woody root systems. METHODS: Five mature, single-stemmed Quercus pubescens trees growing on a steep slope and five on a shallow slope were excavated to a root diameter of 1 cm. A very precise numeric representation of the geometry and topology of structural root architecture was gained using a low-magnetic-field digitizing device (Fastrak, Polhemus). Several characteristics of root architecture were extracted by macros, including root volume, diameter, length, number, spatial position and branching order. KEY RESULTS: The diameter at breast height (dbh) was the best predictor of the root volume but had no correlation with length and number of roots. The slope affected the root volume for each branching order, and the basal cross-sectional area (CSA), number and length of the first-order roots. Number and length of the second- and third-order laterals were closely related in both conditions, although this relationship was closer in the shallow trees, suggesting the influence of a genetic control. Sloping trees showed a clustering tendency of the first- and second-order lateral roots in the up-slope direction, suggesting that the laterals rather than the taproots provide much of the anchorage. In a steep-slope condition, the taproot tapering was positively correlated with the asymmetry magnitude of first-order roots, indicating compensation between taproot and main lateral roots' clustering tendency. CONCLUSIONS: These results suggest that on a slope, on clayey soils, root asymmetry appears to be a consequence of several environmental factors such as inclination, shallow-slides and soil compactness. In addition, this adaptive growth seems to counteract the turning moment induced by the self-loading forces acting in slope conditions, and as a consequence improves the tree stability.     

Development of fine and coarse roots of Thuja occidentalis `Brabant' in non-irrigated and drip irrigated field plots

Aboveground dry mass, total root dry mass and root length density of the fine roots of Thuja occidentalis `Brabant' were determined under non- and drip-irrigated field conditions. Two-dimensional diffusion parameters for dynamic root growth were estimated based on dry mass production of the fine roots and the concept of the convective-diffusion model of cylindrical root growth and proliferation. Drip irrigation increased above-ground dry mass and the shoot:root ratio compared with no irrigation. Dry mass of the coarse roots increased as well due to drip irrigation. No effect on total or fine root dry mass was found. Drip irrigation increased root length densities in the top 0.1 m but not significantly. However, drip irrigation decreased root proliferation in depth by 27%, whereas proliferation in the horizontal direction was not altered. Measured root length densities were overestimated by 6–21% by the model (0.68 < R ² < 0.92).     

黄土丘陵半干旱区密植枣林随树龄变化的根系空间分布特征

该文研究了黄土丘陵半干旱区密植枣( Ziziphus jujuba ‘Lizao’)林群体根系随树龄变化的空间分布特征。对1年生、4年生、8年生和11年生4种树龄的密植枣林采用剖面法, 获得0-1 m土壤剖面上直径>3 mm、1-3 mm及<1 mm的根系数量和空间位置信息。利用方差分析, 评价了山地密植枣林林分根系随树龄变化的水平和垂直分布特征。结果表明: 3种直径的根系数量均随着树龄的增长而增加, 直径< 1 mm的根系增长速度最快; 随着土层加深, 根系数量递减, 1年生枣林的根系主要聚集在0-40 cm土层中, 4年生及以上树龄的根系主要分布在0-60 cm土层中; 0-1 m土层内, 1年生枣林(株距1.2 m)及4年以上树龄(株距2 m), 同树龄枣林中直径<1 mm的根系水平分布无差异; 同一土层中(0-20 cm, 20-40 cm, 40-60 cm), 无论树龄大小及离树干的水平位置如何, 不同直径根系的数量都无差异。研究表明: 在有水肥管理措施的条件下, 枣林根系垂直方向形成浅层型的适应模式; 在密植环境下, 枣林细根形成根网型的适应模式。     

陇中黄土高原主要造林树种细根生物量分布

以2 mm为粗、细根的划分界限,采用根钻法对黄土高原安家沟流域油松、白杨、山杏、刺槐、沙棘和柠条6个主要造林树种细根分布进行研究,并测定不同林地下土壤含水率和土壤理化性质.结果表明： 在水平方向上,油松细根生物量呈先增大后减小的二次多项式分布,其他5个树种细根生物量均呈对数分布,水平根系发达,细根主要分布在冠幅半径2~3倍的范围内,表明各植被通过水平扩展来获取更多的土壤水分.在垂直方向上,随着土层深度的增加,细根生物量均呈减小趋势.6种植被细根生物量与土壤水分、容重呈显著负相关,与有机质、全N含量呈显著正相关.     

Distribution of roots and root nodules and biomass allocation in young intensively managed grey alder stands on a peat bog

Development of below-ground biomass and biomass allocation were studied in two different stands of young grey alder stands growing on a peat bog. Both stands were given the same fertilization and irrigation treatment. The roots were investigated from 1) open plastic tubes enclosing the complete root systems in 1982, and 2) root cores 1984–86. Coarse roots (diameter>1 mm) were mainly found close to the trunk of the trees while fine roots (≤1 mm) were more evenly distributed in the stands. Root nodules were intermediate in distribution. The root systems were shallow, with more than 90% of the biomass in the uppermost 9–10 cm of the soil, probably because of low oxygen availability in the peat soil. The biomass allocation to the above-ground parts increased during the study period.     

The architecture of Picea sitchensis structural root systems on horizontal and sloping terrain

The coarse root systems of 24 Sitka spruce (Picea sitchensis (Bong.) Carr.) trees, from a 40-year-old plantation in west Scotland, were extracted, digitised in three dimensions, and root topology was recorded. Roots were from trees grown on a steep (ca. 30°) north-facing slope, and from an adjacent horizontal area with similar gleyed mineral soil. The prevailing wind was across-slope from the west. Analysis of below-ground parts of the trees in comparison with those above-ground revealed a positive linear relationship between coarse root volume and stem volume. Most non-directional characteristics of the root systems were similar between trees on the slope and on flat terrain. Allocation of root mass around trees was examined in relation to the slope and the prevailing wind direction. Trees on the horizontal area had more root mass in leeward sectors than other sectors, but trees on the slopes had more root mass in the windward sectors than other sectors. Centres of mass of the root systems from the horizontal part of the site were not significantly clustered in any direction, but root systems of trees on the slope had centres of mass significantly clustered across the slope in the windward direction. For trees on the slope, the mean direction of the largest sector without structural roots was 4° from north, i.e. downslope. The results are discussed in relation to soil characteristics and the biomechanical behaviour of trees on slopes.     

Root distribution at various distances from clove trees growing in Indonesia

Efficient fertilizer application requires an understanding of the distribution of roots and soil nutrients in the soil profile. Cultural practices for clove trees in Indonesia has resulted in phosphorus (P) fertilizer being applied at the canopy edge; however, in these high P fixing soils efficient P fertilizer application should occur with the highest root densities. The objective of this study, therefore, was to determine the rooting distribution at various distances from the tree and soil depths for clove (Eugenia aromatica OK; variety Zanzibar) trees growing on an Andosol soil at Modoinding, Indonesia. Root distributions were determined to a 100-cm soil depth using soil cores at 0.5, 1.0 and 1.5 times the canopy radius for five 10-year-old clove trees grown on either level terrain or 23% slopes. Clove root length and weight densities decreased with soil depth and distance from the tree base. Fine clove roots (1 mm dia) comprised 72% of the total root length and was three to five times higher underneath the canopy than that outside the canopy. Roots were concentrated in the upper soil horizons; however, up to 36% of the total root length was found at a depth of 50–100 cm. Clove roots for trees growing at the level landscape position had the highest root length densities. Intercropped species root length densities were higher than clove root length densities at 1.5 times the canopy radius whereas intercropped root weight densities were higher than that for clove roots at both 1.5 and 1 times the canopy radius. Results suggest that fertilizer applications should be placed closer to the tree trunk rather than at the canopy edge to maximize P uptake by clove roots.     

毛竹种群向针阔林扩张的根系形态可塑性

为了弄清毛竹(Phyllostachys edulis)向针阔林扩张过程中根系的形态可塑性反应,在浙江天目山自然保护区毛竹向针阔林扩张的典型过渡地带,连续区域上设置毛竹纯林、针阔-毛竹混交林(以下简称过渡林)、针阔林3种样地。用根钻法采集样地毛竹根系、针阔树根系并比对其生物量密度、细根比根长、相邻同级侧根节点距等形态特征参数变化。结果表明:随着毛竹的扩张程度增加,林内根系生物量密度增加;且与针阔树竞争过程中毛竹将更多的根系放置于表层;同时在水平方向上随离样株距离的增加未出现明显变化,而针阔树根系则随离样木距离的增加而逐渐减少;毛竹根系比根长明显增加,平均增幅15%;一、二级侧根节点距则均有所下降,毛竹侧根数量增多。这些结果表明毛竹种群可通过根系生物量密度、细根比根长、相邻同级侧根节点距等形态可塑性方式实现向周边森林扩张。     

育苗容器规格对橡胶树籽苗芽接苗生长的影响

采用不同规格控根育苗容器对橡胶树籽苗芽接苗进行悬空和近地培养,观测其生长情况。结果表明,悬空培养对橡胶树籽苗芽接苗的纵向伸长(株高、主根长度)影响较为显著。育苗容器的高度一致时,体积越大的育苗容器,根系伸展空间越多,苗木地上部分长势越好;体积一致时,高度越高的育苗容器,苗木主根长度越长,但对籽苗地上部分及主根粗度影响较小。     

Is it possible to manipulate root anchorage in young trees?

The optimal root system architecture for increased tree anchorage has not yet been determined and in particular, the role of the tap root remains elusive. In Maritime pine (Pinus pinaster Ait.), tap roots may play an important role in anchoring young trees, but in adult trees, their growth is often impeded by the presence of a hard pan layer in the soil and the tap root becomes a minor component of tree anchorage. To understand better the role of the tap root in young trees, we grew cuttings (no tap root present) and seedlings where the tap root had (?) or had not (+) been pruned, in the field for 7 years. The force (F) necessary to deflect the stem sideways was then measured and divided by stem cross-sectional area (CSA), giving a parameter analogous to stress during bending. Root systems were extracted and root architecture and wood mechanical properties (density and longitudinal modulus of elasticity, E _L ) determined. In seedlings (?) tap roots, new roots had regenerated where the tap root had been pruned, whereas in cuttings, one or two lateral roots had grown downwards and acted as tap roots. Cuttings had significantly less lateral roots than the other treatments, but those near the soil surface were 14% and 23% thicker than plants (+) and (?) tap roots, respectively. Cuttings were smaller than seedlings, but were not relatively less resistant to stem deflection, probably because the thicker lateral roots compensated for their lower number. Apart from stem volume which was greater in trees (+) tap roots, no significant differences with regard to size or any root system variable were found in plants (?) or (+) tap roots. In all treatments, lateral roots were structurally reinforced through extra growth along the direction of the prevailing wind, which also improved tap root anchorage. Predictors of log F/CSA differed depending on treatment: in trees (?) tap roots, a combination of the predictors stem taper and %volume allocated to deep roots was highly regressed with log F/CSA (R ² = 0.83), unlike plants (+) tap roots where the combined predictors of lateral root number and root depth were best regressed with log F/CSA (R ² = 0.80). In cuttings, no clear relationships between log F/CSA and any parameter could be found. Wood density and E _L did not differ between roots, but did diminish with increasing distance from the stem in lateral roots. E _L was significantly lower in lateral roots from cuttings. Results showed that nursery techniques influence plant development but that the architectural pattern of Maritime pine root systems is stable, developing a sinker root system even when grown from cuttings. Anchorage is affected but the consequences for the long-term are still not known. Numerical modelling may be the only viable method to investigate the function that each root plays in adult tree anchorage.     

Drip irrigation,soil characteristics and the root distribution and root activity of olive trees

A study was carried out on the root distribution and root activity of the olive tree (Olea Europaea, L., var. manzanillo) as influenced by drip irrigation and by several soil characteristics such as texture and depth. The experiments were conducted in two plots within a drip-irrigated grove of 20-year-old trees planted at 7×7 m spacing. One soil was a sandy loam, the other a clay-loam. Both cylinder and trench methods were used to determine root distribution. Labelling with ³²P was used to determine root activity. Under dryland conditions the adult tree adapted its rooting system, following the installation of a drip system, by concentrating the roots within the wet soil zones near the drippers. The highest root densities occur in those zones, down to a 0.6 m depth, the most abundant being the <0.5 mm diameter roots. The most intensive root activity was also found in that zone. For a given irrigation system, wet soil bulbs are more extensive and therefore root distribution expands to a larger soil volume when the soil is more clayey and with a hard calcareous pan present at about 0.8 m depth which prevents deep drainage.     

渭北旱塬不同龄苹果细根空间分布特征

以渭北旱塬3龄、10龄、15龄和20龄苹果树为对象,采用根钻法,沿3等分园半径方向(径向)、距树干1.0、1.5m和2 0m处设置采样点,研究了不同树龄的细根空间分布特征.结果表明,3龄苹果细根主要分布于径向1.5m以内和垂向0.5m以上,15龄和20龄苹果细根分布超出径向2.0m和垂向1.4m,10龄细根分布范围大于3龄,与15龄和20龄接近.在根系主要分布区内3龄和10龄细根分布稀疏,15龄和20龄细根分布密集;细根空间分布演化过程可分为3个阶段,即3～10龄为细根范围扩张阶段,10～15龄为细根密度扩张阶段,15～20龄为细根密度退化阶段;苹果细根空间分布无明显方向性差异;10龄、15龄和20龄苹果表层(0～20cm)平均根长密度低于下层(20～40cm),高峰值一般出现在40～80cm,此深度以下根长密度随深度递减,3龄苹果表层平均根长密度高于下层;在径向2.0m内随径向距离增大,3龄、15龄和20龄平均根长密度逐渐降低,而10龄根长密度逐渐增加.根长密度在径向变化上存在局部变异现象.     

Citrus root responses to localized drying soil: A new approach to studying mycorrhizal effects on the roots of mature trees

Because fine roots tend to be concentrated at the soil surface, exposure to dry surface soil can have a large influence on patterns of root growth, death and respiration. We studied the effects of arbuscular mycorrhizas (AM) formation on specific root length (SRL), respiration and mortality of fine roots of bearing red grapefruit (Citrus paradisi Macf.) trees on Volkamer lemon (C. volkameriana Tan. & Pasq.) rootstock exposed to drying soil. For each tree, the fine roots were removed from two woody lateral roots, the roots were surface sterilized and then each woody root was placed in a separate pair of vertically divided and independently irrigated soil compartments. The two split-pot systems were filled with sterilized soil and one was inoculated with arbuscular mycorrhizal fungi (Glomus etunicatum/G. intraradices). New fine lateral roots that emerged from the woody laterals were permitted to grow inside the pots over a 10-month period. Irrigation was then removed from the top compartment for a 15-week period. At the end of the study, roots inoculated with AM fungi exhibited about 20% incidence of AM formation, whereas the uninoculated roots were completely void of AM fungi. Arbuscular mycorrhizal roots exhibited lower SRL, lower root/soil respiration and about 10% lower fine root mortality than nonmycorrhizal roots after 15 weeks of exposure to dry surface soil. This study demonstrates the feasibility of examining mycorrhizal effects on the fine roots of adult trees in the field using simple inexpensive methods.     

Influence of storage methods on corn root length measurements

This study evaluated the changes in root length, mass, and diameter after air drying and rehydration of corn (Zea mays L.) root samples. For corn roots washed from soil, rehydrated root length was not reduced when compared with fresh root length, but rehydrated root mass was reduced to about half of fresh root mass, and rehydrated root diameter was approximately 75% of fresh diameter. Three storage methods (air dried, 70% ethanol, and 5% formaldehyde solution) were also compared for corn roots grown in moist paper towels. Although root mass and diameter were significantly reduced by air drying, root length was not altered by any of the treatments.     

种间互作和施氮对蚕豆/玉米间作生态系统地上部和地下部生长的影响

为河西走廊绿洲灌区豆科/禾本科间作体系的养分管理提供科学依据,于2007年在武威绿洲农业试验站应用田间原位根系行分隔技术研究了蚕豆/玉米种间互作和施氮对玉米抽雄期的根系空间分布、根系形态和作物地上部生长的影响。研究结果表明:种间互作和施氮均增加了玉米和蚕豆在纵向和横向两个尺度上的根重密度、根长密度、根表面积、根系体积。根长密度和根表面积与两种作物产量和氮素吸收均呈正相关,而与蚕豆的根瘤重呈负相关;抽雄期的土壤含水量与玉米产量和养分吸收呈显著的负相关。玉米根系可以占据蚕豆地下部空间,但蚕豆的根却较少到间作玉米的地下部空间,也就是间作后增加了玉米根系水平尺度的生态位。蚕豆和玉米根系主要分布分别在0-40 cm浅土层和0-60 cm 土层,且间作玉米根系在60-120 cm比单作和分隔的多。因此,种间互作和施氮扩大了两作物根系纵向和横向的空间生态位,改变了作物根系形态,即扩展了两者水分和养分吸收的生态位,增加了作物吸收养分的有效空间,从而提高了间作生态系统的生产力。