Nitrogen uptake and transformation in a midwestern U.S. stream: A stable isotope enrichment study

This study presents a comprehensive analysis ofnitrogen (N) cycling in a second-order forestedstream in southern Michigan that has moderatelyhigh concentrations of ammonium (mean,16 g N/L) and nitrate (17 g N/L). Awhole-stream ¹⁵NH₄ ⁺ addition wasperformed for 6 weeks in June and July, and thetracer ¹⁵N was measured downstream inammonium, nitrate, and detrital and livingbiomass. Ancillary measurements includedbiomass of organic matter, algae, bacteria andfungi, nutrient concentrations, hydrauliccharacteristics, whole-stream metabolism, andnutrient limitation assays. The resultsprovide insights into the heterotrophic natureof woodland streams and reveal the rates atwhich biological processes alter nitrogentransport through stream systems.Ammonium uptake lengths were 766–1349 m anduptake rates were 41–60 g N m^–2min^–1. Nitrate uptake could not bedetected. Nitrification rates were estimatedfrom the downstream increase in¹⁵N-enriched nitrate using a simulationmodel. The ammonium was removed bynitrification (57% of total uptake),heterotrophic bacteria and fungi associatedwith detritus (29%), and epilithic algae(14%). Growth of algae was likely limited bylight rather than nutrients, and dissolvedO₂ revealed that the stream metabolism washeterotrophic overall (P:R = 0.2). Incubationsof detritus in darkened chambers showed thatuptake of ¹⁵N was mostly heterotrophic.Microbial N in detritus and algal N inepilithon appeared to reach isotopic steadystate with the dissolved ammonium, but theisotopic enrichment of the bulk detritus andepilithon did not approach that of ammonium,probably due to a large fraction of organic Nin the bulk samples that was not turning over. The actively cycling fraction of total N inorganic compartments was estimated from theisotopic enrichment, assuming uptake ofammonium but not nitrate, to be 23% forepilithon, 1% for fine benthic organic matter,5% for small woody debris, and 7% for leaves. These percentages agree with independentestimates of epilithic algal biomass, whichwere based on carbon:chlorophyll ratios in bulksamples and in algal fractions separated bydensity-gradient centrifugation in colloidalsilica, and of microbial N in the detritus,which were based on N released by chloroformfumigations.     

Nitrogen Transfer Within and Between Plants Through Common Mycorrhizal Networks (CMNs)

Mycorrhizae play a critical role in nutrient capture from soils. Arbuscular mycorrhizae (AM) and ectomycorrhizae (EM) are the most important mycorrhizae in agricultural and natural ecosystems. AM and EM fungi use inorganic NH₄ ⁺ and NO₃ ^?, and most EM fungi are capable of using organic nitrogen. The heavier stable isotope ¹⁵N is discriminated against during biogeochemical and biochemical processes. Differences in ¹⁵N (atom%) or δ¹⁵N (‰) provide nitrogen movement information in an experimental system. A range of 20 to 50% of one-way N-transfer has been observed from legumes to nonlegumes. Mycorrhizal fungal mycelia can extend from one plant's roots to another plant's roots to form common mycorrhizal networks (CMNs). Individual species, genera, even families of plants can be interconnected by CMNs. They are capable of facilitating nutrient uptake and flux. Nutrients such as carbon, nitrogen and phosphorus and other elements may then move via either AM or EM networks from plant to plant. Both ¹⁵N labeling and ¹⁵N natural abundance techniques have been employed to trace N movement between plants interconnected by AM or EM networks. Fine mesh (25～45 μm) has been used to separate root systems and allow only hyphal penetration and linkages but no root contact between plants. In many studies, nitrogen from N₂-fixing mycorrhizal plants transferred to non-N₂–fixing mycorrhizal plants (one-way N-transfer). In a few studies, N is also transferred from non-N₂–fixing mycorrhizal plants to N₂-fixing mycorrhizal plants (two-way N-transfer). There is controversy about whether N-transfer is direct through CMNs, or indirect through the soil. The lack of convincing data underlines the need for creative, careful experimental manipulations. Nitrogen is crucial to productivity in most terrestrial ecosystems, and there are potential benefits of management in soil-plant systems to enhance N-transfer. Thus, two-way N-transfer warrants further investigation with many species and under field conditions.     

Nitrogen transfer from arrowleaf clover to ryegrass in field plantings

Arrowleaf clover (Trifolium vesiculosum Savi) and annual ryegrass Lolium multiflorum Lam.) commonly are overseeded in dormant bermudagrass (Cynodon dactylon L. Pers.) sod on coastal plain soils in the southeastern United States. Two field experiments were conducted in consecutive years at different sites to estimate the amount of N transferred from the clover to the annual grass. Nitrogen treatments included 50 kg N ha^-1 as ¹⁵N depleted ammonium nitrate applied in either February or April, and a check (no N applied). Three clippings were made during the cool-season from March to June. In both experiments, less than 5 kg N ha^-1 were transferred from the clover to the grass. Ryegrass yields of dry matter and total N were not increased by growing with clover. Clover growth was typical for the region; average dry matter yield in pure stand was 2,615 kg ha^-1 over the two-year period. Clover in mixed stand fixed between 20 and 60 kg N/ha. Less than 13% of N contained in ryegrass was transferred from arrowleaf clover to ryegrass at any clipping while clover was actively growing. The quantity of N transferred over the entire season was not statistically significant.     

Nitrogen assimilation in field-grown winter wheat: Direct measurements of nitrate reduction in roots using15N

Summary Nitrate fertiliser labelled with¹⁵N was applied to a field grown crop of winter wheat. Uptake and assimilation of fertiliser nitrate was studied by monitoring the appearance of labelled nitrate and labelled amino acids in the xylem sap. Shortly after applying¹⁵N-nitrate to the soil about 30 per cent of recently absorbed¹⁵N was in the reduced form, indicating that roots of cereal crops can make a substantial contribution in reducing nitrate. Seasonal changes in crop growth andin vivo NRA are also described.     

Introgressive hybridization between Galaxias depressiceps and Galaxias sp D (Teleostei: Galaxiidae) in Otago, New Zealand: Secondary contact mediated by water races

Central Otago (New Zealand) was the focus of an intense period ofgold mining in the late 19th century. Mining relied on high waterpressure for sluicing, often requiring extensive systems of water raceslinking headwaters over large distances. On Rough Ridge, races connectedheadwaters of Otago's two major river systems, the Taieri and theClutha. We employed genetic analysis of freshwater fish(Galaxias) populations to assess the biological implications ofrace-development. The two galaxiid taxa found on Rough Ridge (G. spD and G. depressiceps) are reciprocally monophyletic formtDNA and have three fixed allozyme differences. Rough Ridge aside,G. sp D is apparently absent from the Taieri system and G.depressiceps from the Clutha system. Distributions of mitochondrialDNA, allozyme and microsatellite alleles imply that artificialconnections have facilitated bidirectional introgression: movementof G. sp D alleles into the Taieri system and G.depressiceps alleles into the Clutha. The finding of some identicalcontrol region haplotypes (of both lineages) on either side of thecatchment boundary confirms the recency of genetic exchange. Near theinferred region of contact, samples from Linn Burn (Taieri) are pureG. depressiceps for three diagnostic allozymes and mtDNA.Samples from upper Pool Burn (Clutha) are pure G. sp D forthese same four markers. However, other collections from around thecontact zone represent genetic mixtures forming a cline. We fear thatsimilar human-induced mixing may have happened elsewhere in uplandOtago.     

Nitrogen fixation in breeding lines of Phaseolus vulgaris L.

Nitrogen fixation in nine common bean (Phaseolus vulgaris L.) lines was estimated using the ¹⁵N isotope dilution method at two locations in two seasons. In the first season at one location no N₂ fixation was detected while in the second season up to 51 kg N ha^–1 were estimated. There were significant differences between lines and correlations between trials were significant for the amounts of N₂ fixed, but not for total shoot nitrogen. The plants that fixed the most nitrogen nodulated rapidly after germination. Differences in maximum nodule mass, but not specific nodule activity, were detected also.     

基于δ15N稳定同位素分析的人工防护林大型土壤动物营养级研究

稳定同位素δ15N是分析生态系统中的营养关系的重要方法。本研究采用该方法分析了人工防护林土壤动物的营养级及它们之间的营养关系。对3个地区人工杨树林12类大型土壤动物及其土壤、细根及植物枯落叶的稳定同位素δ15N分析结果显示,不同环境条件下土壤动物δ15N具有显著差异性(F=38.067,P0.001),土壤动物δ15N值与土壤、植物细根及枯落叶δ15N值之间有明显的相关性,与全氮含量没有明显的相关性;根据不同土壤动物δ15N值的状况,12种土壤动物在土壤生态系统中大都处于第二至第四营养级,处于较低营养级位置的动物主要有大蚊幼虫(Tipulidae larvae)、蚯蚓(Lumbricidae)、金龟甲幼虫(Scarabaeidae larvae)等类群,较高营养级的为蜈蚣(Scolopendromorpha)、隐翅甲成虫(Staphylinidae adult)、叩甲成虫(Elateridae adult)、线蚓(Enchytraeidae)等动物,同类土壤动物在不同样地中营养级位置存在一定的差异。     

不同氮处理下速生柳对水体氮的吸收、分配及生理响应

采用水培法,研究了旱柳苗在外源添加不同氮水平(贫氮、中氮、富氮、过氮)的铵态氮(NH+4-N)和硝态氮(NO-3-N)的生长、氮吸收、分配和生理响应。结果表明:一定范围氮浓度的增加能够促进旱柳苗的生长,但过量氮会抑制其生长,且NH+4-N的抑制作用大于NO-3-N;两种氮处理下,旱柳表现出对NH+4-N的吸收偏好,在同一氮水平时,旱柳各部位氮原子百分含量Atom%15N(AT%)、15N吸收量和来自氮源的N%(Ndff%)均为NH+4-N处理大于NO-3-N处理,且随着氮浓度的增加,差异增大,且在旱柳各部位的分布为根﹥茎﹥叶;2种氮素过量和不足均会对旱柳根和叶生理指标产生不同的影响,其中在过氮水平时,NH+4-N和NO-3-N处理下根系活力比对照减少了50.61%和增加了19.53%;在过氮水平时,NH+4-N处理柳树苗根总长、根表面积、根平均直径、根体积和侧根数分别对照下降了30.92%、29.48%、19.44%、27.01%和36.41%,NO-3-N处理柳树苗相应的根系形态指标分别对对照下降了1.66%、5.65%、1.49%、5.06%和25.72%。可见,高浓度NH+4-N对旱柳苗的胁迫影响大于NO-3-N,在应用于水体氮污染修复时可通过改变水体无机氮的比例,削弱其对旱柳的影响,从而提高旱柳对水体氮污染的修复效果。     

Changes in Canopy Processes Following Whole-Forest Canopy Nitrogen Fertilization of a Mature Spruce-Hemlock Forest

Abstract Most experimental additions of nitrogen to forest ecosystems apply the N to the forest floor, bypassing important processes taking place in the canopy, including canopy retention of N and/or conversion of N from one form to another. To quantify these processes, we carried out a large-scale experiment and determined the fate of nitrogen applied directly to a mature coniferous forest canopy in central Maine (18–20 kg N ha⁻¹ y⁻¹ as NH₄NO₃ applied as a mist using a helicopter). In 2003 and 2004 we measured NO₃ ⁻, NH₄ ⁺, and total dissolved N (TDN) in canopy throughfall (TF) and stemflow (SF) events after each of two growing season applications. Dissolved organic N (DON) was greater than 80% of the TDN under ambient inputs; however NO₃ ⁻ accounted for more than 50% of TF N in the treated plots, followed by NH₄ ⁺ (35%) and DON (15%). Although NO₃ ⁻ was slightly more efficiently retained by the canopy under ambient inputs, canopy retention of NH₄ ⁺as a percent of inputs increased markedly under fertilization. Recovery of less than 30% of the fertilizer N in TF suggested that the forest canopy retained more than 70% of the applied N (>80% when corrected for N which bypassed tree surfaces at the time of fertilizer addition). Results from plots receiving ¹⁵N enriched NO₃ ⁻ and NH₄ ⁺ confirmed bulk N estimations that more NO₃ ⁻ than NH₄ ⁺ was washed from the canopy by wet deposition. The isotope data did not show evidence of canopy nitrification, as has been reported in other spruce forests receiving much higher N inputs. Conversions of fertilizer-N to DON were observed in TF for both ¹⁵NH₄ ⁺ and ¹⁵NO₃ ⁻ additions, and occurred within days of the application. Subsequent rain events were not significantly enriched in ¹⁵N, suggesting that canopy DON formation was a rapid process related to recent N inputs to the canopy. We speculate that DON may arise from lichen and/or microbial N cycling rather than assimilation and re-release by tree tissues in this forest. Canopy retention of experimentally added N may meet and exceed calculated annual forest tree demand, although we do not know what fraction of retained N was actually physiologically assimilated by the plants. The observed retention and transformation of DIN within the canopy demonstrate that the fate and ecosystem consequences of N inputs from atmospheric deposition are likely influenced by forest canopy processes, which should be considered in N addition studies. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

植被恢复过程中芒萁覆盖对侵蚀红壤氮组分的影响

氮素是限制陆地生态系统生产力的重要因子。采用时空代换法,以红壤侵蚀区未治理、恢复12年和30年的马尾松林为研究对象,对比分析了林下芒萁覆盖地与裸地表层土壤之间氮同位素、不同形态氮组分含量以及不同组分氮含量所占比例之间的差异。结果表明:在所有马尾松林中,芒萁覆盖增加了表层土壤的全氮含量,δ~(15)N值则比林下裸地显著降低了33. 8%—83.1%(P0.05)。随着恢复年限增加,林下芒萁覆盖地表层土壤δ~(15)N值显著下降,而林下裸露地δ~(15)N值没有显著变化(P0.05)。不同恢复年限马尾松林的芒萁覆盖地表层土壤微生物生物量氮、可溶性有机氮和铵态氮含量显著高于林下裸地(P 0.05),而硝态氮含量则显著低于林下裸地(P0.05)。随恢复年限增加,表层土壤微生物生物量氮、可溶性有机氮、铵态氮含量均呈增加趋势,而硝态氮含量则呈下降趋势,不同形态氮占全氮比例表现为:微生物生物量氮铵态氮可溶性有机氮硝态氮。相关分析表明土壤δ~(15)N值与硝态氮极显著正相关,与其他氮组分极显著负相关(P0.01)。由此可见,与林下裸地相比,芒萁覆盖在植被恢复过程中有助于提高表层土壤中全氮、微生物生物量氮、可溶性有机氮和铵态氮含量,降低硝态氮的淋溶损失风险,促进土壤氮保持和积累,从而有利于退化红壤生态系统的恢复。     

Nitrogen uptake from prey and substrate as affected by prey capture level and plant reproductive status in four carnivorous plant species

Uptake of nitrogen from prey and substrate and partitioning of prey-derived nitrogen were studied in the carnivorous plant species Pinguicula alpina, P. villosa, P. vulgaris and Drosera rotundifolia in a subarctic environment. Efficiency in nitrogen uptake from prey was evaluated by tracing ¹⁵N from ¹⁵N-enriched Drosophila flies fed to the plants. The in situ uptake efficiency differed somewhat between species and ranged from 29 to 41% of prey N. This efficiency was not affected by different feeding levels or plant reproductive status (flowering or non-flowering). A test of the amount of N absorbed from prey caught on flower stalks of Pinguicula villosa and P. vulgaris showed that both species took up little of what was available in prey (2.5% or less). The uptake efficiency found in greenhouse grown plants was higher than in plants in situ (40–50% vs. 30–40% respectively). This could probably best be explained by the absence of rain and a higher temperature in the greenhouse. The prey-derived ¹⁵N was traced to reproductive organs and winter buds. Non-flowering individuals allocated 58–97% of the N derived from prey to their winter buds. Flowering individuals allocated 17–43% of the N income from prey to reproduction, while 34–71% were allocated to buds. Root uptake of nitrogen was stimulated by increased prey capture. This increase in uptake of nitrogen from the substrate was larger than the potential direct uptake of nitrogen from captured prey.     

Carbon and nitrogen turnover in adjacent grassland and cropland ecosystems

The effects of cultivation and soil texture on net and gross N mineralization, CO₂ evolution and C and N turnover were investigated using paired grassland and cropped sites on soils of three textures. Gross N mineralization and immobilization were measured using¹⁵N-isotope dilution. Grassland soils had high CO₂ evolution and gross N mineralization rates, and low net N mineralization rates. Cropland soils had low CO₂ evolution rates but had high net and gross N mineralization rates. Grassland soils thus had high immobilization rates and cropland soils had low immobilization rates. Cultivation increased N turnover but reduced C turnover. The data suggest that the microflora in grassland soils are N limited, while those of cropland soils are limited by C availability. Increasing clay content reduced N turnover. C turnover was less clearly related to texture. Differences in the immobilization potential of substrates help explain why agricultural soils have higher N losses than do grassland soils.     

Fate of urine nitrogen on mineral and peat soils in New Zealand

A field lysimeter experiment was conducted over 150 days to examine the fate of synthetic urinary nitrogen (N) applied to peat and mineral soils, with and without a water table. At the start of the winter season, synthetic urine labelled with ¹⁵N, was applied at 500 kg N ha^–1. Plant uptake, leaching losses and nitrous oxide (N₂O) fluxes were monitored. Total plant uptake ranged from 11% to 35% of the urine-N applied depending on soil type and treatment. Plant uptake of applied N was greater in the presence of a water table in the mineral soil. Nitrate-N (NO₃ ^--N) was only detected in leachates from the mineral soil, at concentrations up to 146 g NO₃ ^--N mL^–1. Presence of a water table in the mineral soil reduced leaching losses (as inorganic-N) from 47% to 6%, incrased plant uptake and doubled apparent denitrification losses. In the peat soils leaching losses of applied urine-N as inorganic-N were low (<5%). Losses of N as N₂O were greater in the mineral soil than in the peat soils, with losses of 3% and <1% of N applied respectively after 100 days. Apparent denitrification losses far exceeded N₂O losses and it is postulated that the difference could be due to dinitrogen (N₂) loss and soil entrapment of N₂.     

Nitrogen fixation and metabolism by groundwater-dependent perennial plants in a hyperarid desert

The Central Asian Taklamakan desert is characterized by a hyperarid climate with less than 50 mm annual precipitation but a permanent shallow groundwater table. The perched groundwater (2–16 m) could present a reliable and constant source of nitrogen throughout the growing season and help overcome temporal nitrogen limitations that are common in arid environments. We investigated the importance of groundwater and nitrogen fixation in the nitrogen metabolism of desert plants by assessing the possible forms and availability of soil N and atmospheric N and the seasonal variation in concentration as well as isotopic composition of plant N. Water availability was experimentally modified in the desert foreland through simulated flooding to estimate the contribution of surface water and temporally increased soil moisture for nutrient uptake and plant–water relations. The natural vegetation of the Taklamakan desert is dominated by plants with high foliar nitrogen concentrations (2–3% DM) and leaf nitrate reductase activity (NRA) (0.2–1 mol NO₂^– g^–1 FW h^–1). There is little evidence that nitrogen is a limiting resource as all perennial plants exhibited fast rates of growth. The extremely dry soil conditions preclude all but minor contributions of soil N to total plant N so that groundwater is suggested as the dominant source of N with concentrations of 100 M NO₃^–. Flood irrigation had little beneficial effect on nitrogen metabolism and growth, further confirming the dependence on groundwater. Nitrogen fixation was determined by the ¹⁵N natural abundance method and was a significant component of the N-requirement of the legume Alhagi, the average contribution of biologically fixed nitrogen in Alhagi was 54.8%. But nitrogen fixing plants had little ecological advantage owing to the more or less constant supply of N available from groundwater. From our data we conclude that the perennial species investigated have adapted to the environmental conditions through development of root systems that access groundwater to satisfy demands for both water and nutrients. This is an ecologically favourable strategy since only groundwater is a predictable and stable resource.     

Nitrogen fixation by soybeans (Glycine max L.) in Zambia

Soybeans (Glycine max L.) are being introduced as a cash crop to small scale farmers in Zambia for rotation in their farming systems. The objectives of this study were to compare and select the most approriate non-fixing reference crop for estimating N₂ fixation by soybeans and assess yields and N₂ fixation of soybeans in Zambia. Nitrogen isotope dilution techniques using¹⁵N-labelled organic or inorganic materials were utilized. Two nonnodulating soybean cultivars, Clark RJ1 and N77 or in their absence Pearl millet (Panicum glaucum L.) were judged to be appropriate reference crops. A local soybean fixing cultivar (Glycine max L. cv. Magoye) rated highest among three cultivars tested for its ability to support symbiotic N₂ fixation byB. japonicum under the experimental conditions. Values of percent N derived from atomosphere for this cultivar were in the order of 65 to 70%.deceased.Contribution no R531 of the Saskatchewan Institute of Pedology. Present address (REK): Esso Chemical Canada, P.O. Box 3010, Lethbridge, Alberta Canada T1J 4A9.     

干旱胁迫下胡杨实生幼苗氮素吸收分配与利用

胡杨(Populus euphratica)是塔里木河流域荒漠河岸林的建群种,水分和氮素是限制胡杨幼苗的存活及早期生长的主要因子。利用~(15)N同位素示踪技术分析水和氮素的交互作用对胡杨幼苗不同生长阶段氮素的吸收分配利用及幼苗生长的影响,进一步探究氮素对胡杨实生苗早期形态建成的作用及对干旱胁迫的缓解效应,以期提高幼苗的存活率。实验以一年生胡杨实生幼苗为研究对象,采用温室内盆栽实验,设置4个干旱处理(D_1、D_2、D_3、D_4,土壤相对含水量为:20%—25%、40%—45%、60%—65%、80%—85%)和3种氮素水平(N_0、N_1、N_2:0、3、6 g/盆)测定胡杨幼苗的生长指标和各部分的Ndff、分配率及利用率。结果表明:胡杨幼苗在土壤相对含水量60%—65%(D_3)、氮素添加量3 g/盆(N_1)时,其生长表现为最佳状态;干旱胁迫下,不同氮素添加量对胡杨幼苗各部分的Ndff值存在显著差异,N_2低于N_1;随干旱胁迫减弱(D_3、D_4),植株在生长早期(25 d)根部吸收的~(15)N优先向地上部分转运,生长后期(75 d)植株Ndff最高,其中以根系中Ndff最高;不同生长期幼苗各部分的~(15)N分配存在显著差异,根系~(15)N分配率较高,但不同氮量处理间差异不显著;随生长期的推移,植株对~(15)NH_4~(15)NO_3的利用率表现为粗根最大,各处理中D_3N_1处理均显著高于其他处理。结论:轻度干旱胁迫下添加适量的氮素能够增强植株对氮素的吸收征调能力,优化水资源获取以维持生存的重要机制。     

Nitrogen uptake in needles of Scots pine (Pinus sylvestris L.) when exposed to gaseous ammonia and ammonium fertilizer in the soil

Young saplings of Pinus sylvestris L. were exposed to gaseous NH₃ at 53 or 105 g m^–3 for one year in open-top chambers. Saplings received ¹⁵N-labelled (NH₄)₂SO₄ via the soil. To examine the importance of foliar N uptake, changes in the concentration of total and labelled N in the needles were followed. Increase in needle biomass and N concentration were found in trees exposed to NH₃, confirming that atmospheric NH₃ acted as a N fertilizer. NH₃ had a greater and quicker effect than (NH₄)₂SO₄: compared with the growth in ambient air, the N concentration in the needles exposed to NH₃ had increased by 49% in four months, while the increase after highest N-fertilization (200 kg N ha^–1 y^–1) was only 8%. The small contribution of NH₄ ⁺ fertilization to the total N concentration was not due to a deficient N uptake: the ¹⁵N concentration in the needles increased significantly with time. On the other hand, NH₃ uptake in shoots may have a negative effect on the NH₄ ⁺ root uptake. The relation between plant N and atmospheric NH₃ concentration was non-linear and possible reasons for this observation are discussed. Fumigation with NH₃ significantly decreased the ratios of K/N and P/N, showing that fumigation disrupted the nutrient balance.     

Effects of earthworms on biomass production,nitrogen allocation and nitrogen transfer in wheat-clover intercropping model systems

The effects of earthworms (Lumbricidae) on plant biomass production and N allocation in model intercropping systems of winter wheat and white clover were evaluated in two pot experiments. Wheat and wheat-clover mixtures were grown in a low-organic loam soil, earthworms were added at densities comparable to field population densities and the experiments were terminated 48 and 17 d after earthworm introductions. In both experiments, earthworms significantly increased the biomass and N uptake of wheat while they had generally no effects on clover. As a result, earthworm activity increased the proportion of wheat biomass in the total plant biomass of the mixture. Nitrogen budgets of the experiment lasting 48 d indicated that additional N in the system made available by earthworm activity was primarily taken up by the wheat. Earthworms also affected intra-plant N allocation in wheat which had significantly higher shoot:root N ratios when earthworms were present. When clover was labelled with ¹⁵N in the experiment which lasted 17 d, endogeic earthworms significantly reduced the amounts of ¹⁵N excess transferred from living or decomposing clover roots to accompanying wheat plants. Earthworms assimilated small quantities of ¹⁵N tracer from decomposing clover roots but not from living clover roots. The results of these model experiments suggest that earthworms can affect the balance between intercropped cereals and legumes by altering intra- and inter-plant N allocation. This revised version was published online in June 2006 with corrections to the Cover Date.