Bi-directional transfer of nitrogen between alfalfa and bromegrass: Short and long term evidence

Transfer of N from legumes to associated non-legumes has been demonstrated under a wide range of conditions. Because legumes are able to derive their N requirements from N₂ fixation, legumes can serve, through the transfer of N, as a source of N for accompanying non-legumes. Studies, therefore, are often limited to the transfer of N from the legume to the non-legume. However, legumes preferentially rely on available soil N as their source of N. To determine whether N can be transferred from a non-legume to a legume, two greenhouse experiments were conducted. In the short-term N-transfer experiment, a portion of the foliage of meadow bromegrass (Bromus riparius Rhem.) or alfalfa (Medicago sativa L.) was immersed in a highly labelled ¹⁵N-solution and following a 64 h incubation, the roots and leaves of the associated alfalfa and bromegrass were analyzed for ¹⁵N. In the long-term N transfer experiment, alfalfa and bromegrass were grown in an ¹⁵N-labelled nutrient solution and transplanted in pots with unlabelled bromegrass and alfalfa plants. Plants were harvested at 50 and 79 d after transplanting and analyzed for ¹⁵N content. Whether alfalfa or bromegrass were the donor plants in the short-term experiment, roots and leaves of all neighbouring alfalfa and bromegrass plants were enriched with ¹⁵N. Similarly, when alfalfa or bromegrass was labelled in the long-term experiment, the roots and shoots of neighbouring alfalfa and bromegrass plants became enriched with ¹⁵N. These two studies conclusively show that within a short period of time, N is transferred from both the N₂-fixing legume to the associated non-legume and also from the non-legume to the N₂-fixing legume. The occurrence of a bi-directional N transfer between N₂-fixing and non-N₂-fixing plants should be taken into consideration when the intensity of N cycling and the directional flow of N in pastures and natural ecosystems are investigated.     

A comparison between the acetylene reduction method,the isotope dilution method and the total nitrogen difference method for measuring nitrogen fixation in lucerne (Medicago sativa L.)

Summary In view of the increasing need for the exact estimation of the input of nitrogen in agroecosystems, an application of the acetylene-reduction technique was developed. The technique, consisting of a plastic bag incubation system, using propane as an internal standard of the apparent volume, made it possible to carry out repeated incubations on the same plant system. The evaluated technique included studies of the diffusion of ethylene and propane in a soil column, as well as studies of the optimal substrate concentration and grade of purity of the substrate. In addition, the conversion factor between amounts of reduced acetylene as compared wtih reduced dinitrogen was determined by¹⁵N₂ incubations to be 4.41. The developedin situ acetylene-reduction technique was compared with an isotope dilution method and a total nitrogen difference method. By comparing the derived total nitrogen fixation values from each with the value derived from the acetylene-reduction method; it was shown that the values differed significantly. The acetylene-reduction method gave the highest nitrogen fixation values, the isotope dilution the lowest values and the total nitrogen difference method was intermediate. No statistical significant difference existed between the two different reference crops used in the isotope dilution method.     

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.     

Nitrate deposition in northern hardwood forests and the nitrogen metabolism of Acer saccharum marsh

It is generally assumed that plant assimilation constitutes the major sink for anthropogenic Nitrate NO ₃ ^– deposited in temperate forests because plant growth is usually limited by nitrogen (N) availability. Nevertheless, plants are known to vary widely in their capacity for NO ₃ ^– uptake and assimilation, and few studies have directly measured these parameters for overstory trees. Using a combination of field and greenhouse experiments, we studied the N nutrition of Acer saccharum Marsh. in four northern hardwood forests receiving experimental NO ₃ ^– additions equivalent to 30 kg N ha^–1 year^–1. We measured leaf and fine-root nitrate reductase activity (NRA) of overstory trees using an in vivo assay and used ¹⁵N to determine the kinetic parameters of NO ₃ ^– uptake by excised fine roots. In two greenhouse experiments, we measured leaf and root NRA in A. saccharum seedlings fertilized with 0–3.5 g NO ₃ ^– –N m^–2 and determined the kinetic parameters of NO ₃ ^– and NH ₄ ⁺ uptake in excised roots of seedlings. In both overstory trees and seedlings, rates of leaf and fine root NRA were substantially lower than previously reported rates for most woody plants and showed no response to NO ₃ ^– fertilization (range = non-detectable to 33 nmol NO ₂ ^– g^–1 h^–1). Maximal rates of NO ₃ ^– uptake in overstory trees also were low, ranging from 0.2 to 1.0 mol g^–1 h^–1. In seedlings, the mean V _max for NO ₃ ^– uptake in fine roots (1 mol g^–1 h^–1) was approximately 30 times lower than the V _max for NH ₄ ⁺ uptake (33 mol g^–1 h^–1). Our results suggest that A. saccharum satisfies its N demand through rapid NH ₄ ⁺ uptake and may have a limited capacity to serve as a direct sink for atmospheric additions of NO ₃ ^– .     

Remobilization of labelled nitrogen from vegetative tissues to pods of mungbean

Summary Remobilization of¹⁵N from vegetative tissue of mungbean (Vigna radiata (L.) Wilczek) into pods was measured during the reproductive phase of growth. Plant tissue was labelled with¹⁵N during vegetative development. Experiments were conducted in the field at two sites. At one site the soil provided cowpeas with most of their N but at the other site N fixation provided most of the N. Remobilized N from vegetative tissue to pods occurred soon after they began to develop. The quantity of the labelled N ultimately remobilized to the pods amounted to 50% for one cultivar (Tx33) at the high soil N site and 70% at the low N site. For the other cultivar (Tx13) the values were 25% and 30%, respectively. The two cultivars performed very differently with respect to partitioning of N into pods and the rate of N fixation. Even though more N was accumulated in the shoots of the high N fixing cultivar (Tx13) less total N was contained in the pods.     

Effect of white clover cultivar on apparent transfer of nitrogen from clover to grass and estimation of relative turnover rates of nitrogen in roots

The apparent transfer of N from clover to associated grass was evaluated over a four year period both on the basis of harvested herbage and by taking account of changes in N in stubble and root (to 10 cm depth) in swards with perennial ryegrass and three different white clover cultivars differing in leaf size. The large leaved Aran transferred 15% of its nitrogen while Huia transferred 24% and the small leaved Kent Wild White transferred 34%. When changes in stubble and root N were taken into account the percentage of N transferred was calculated to be 5% less than in harvested herbage only, as the small leaved types had proportionately more N in the roots and stolons, but the large leaved type was probably more competitive towards the grass.Loss of N from clover roots from July to October was compared to that from grass roots in a grass/white clover sward continuously stocked with steers using a method which incorporated tissue turnover and ¹⁵N dilution techniques. Less than 1 mg N m^-2 d^-1 was lost from the grass roots. In contrast 8 mg m^-2 d^-1 were estimated to be lost from clover roots while 12 mg N m^-2 d^-1 were assimilated.It is concluded that clover cultivar and competitive ability on grass have to be taken into account together with the relationship between N turnover in roots and N available for grass growth when modelling N transfer in grass/clover associations.     

Below-ground nitrogen transfer between different grassland species: Direct quantification by 15N leaf feeding compared with indirect dilution of soil 15N

Nitrogen (N) transfer from one species to another is important for the N cycling in low-input grassland. In the present work, estimates obtained by an indirect ¹⁵N dilution technique were compared with estimates obtained by a direct ¹⁵N leaf feeding technique over two complete growing seasons in red clover-ryegrass and white clover-ryegrass mixtures under field conditions. The direct technique confirmed that N transfer between clovers and ryegrass is a bi-directional process. The transfer of N from both clovers to ryegrass occurred within 25 days upon the first labelling event. A very high N transfer occurred from white clover to the associated ryegrass, 4.5 and 7.5 g m⁻² in the 1st and 2nd production year, respectively. The corresponding values for transfer from red clover to the associated ryegrass were 1.7 and 3.6 g m⁻². Quantified relatively to the total above-ground N content of white clover- ryegrass and red clover-ryegrass mixtures, the N transfer exceeded 50% and 10%, respectively, in three out of seven harvests. The N transfer from ¹⁵N labelled grass to associated clovers constituted a relatively constant proportion of approx. 8% of the above-ground N content of the mixtures. Estimates based on the soil ¹⁵N dilution technique generally underestimated the net N transfer by more than 50% compared to the direct ¹⁵N labelling technique. Furthermore, the indirect ¹⁵N dilution technique estimated only marginal differences between red and white clover in the quantities of N transferred, whereas the direct ¹⁵N labelling technique showed the N transfer from white clover to the associated ryegrass to be significantly higher than that involving red clover. It is concluded that N transfer is a much more dynamic and quantitatively important process in grassland than previously recognised. This revised version was published online in June 2006 with corrections to the Cover Date.     

Variation in the natural abundance of 15N in the halophyte,Salicornia virginica,associated with groundwater subsidies of nitrogen in a southern California salt-marsh

To provide insight into the importance of the salt-marsh ecotone as a sink for inorganic nitrogen in perched groundwater, measurements were made of the natural abundance of ¹⁵N in dissolved NO₃-N and NH₄-N and in the salt-marsh halophyte, Salicornia virginica, along an environmental gradient from agricultural land into a salt-marsh. The increase in the natural abundance of ¹⁵N (expressed by convention as ¹⁵N) of NO₃-N, accompanied by the decrease in NO₃-N (and total dissolved inorganic N, DIN) concentration along the gradient, suggested that the salt-marsh ecotone is a site of transformation, most likely through denitrification, of inorganic nitrogen in groundwater. ¹⁵N enrichment in S. virginica (and the parasitic herb, Cuscuta salina), along the tidal marsh boundary, relative to high and middle marsh locations, indicated the retention of groundwater nitrogen as vegetative biomass. The correlation between ¹⁵N _Salicornia and ¹⁵N_NH4 suggested a preference for NH₄-N over NO₃-N during uptake by this plant. Groundwater inputs enhanced the standing crop, above-ground productivity, and nitrogen content of S. virginica but the ralative effects of pore water salinity and DIN concentration on these parameters were not determined. ¹⁵N enrichment of marsh plants by groundwater DIN inputs could prove useful in tracing the fate of these inputs in the marsh food web.     

A further extension of the method for independently measuring the amount of nitrogen fixed by a legume crop

Summary The combination of using¹⁵N for determining the amount of nitrogen fixed by a legume crop in field experiments and the labelling of only one treatment at a time in each treatment combination is shown to be conceptually and experimentally valid for determining the effect of cultural practices on the amount of nitrogen fixed by a legume crop.     

Forests losing large quantities of nitrogen have elevated 15N:14N ratios

Summary Urea (U) and ammonium nitrate (AN) had been applied to a Scots pine (Pinus sylvestris L.) forest in northern Sweden for 18 consecutive years at four doses resulting in total N applications ranging from 0 to 1980 kg ha^–1. The ¹⁵N abundance ( ¹⁵N) of the grass Deschampsia flexuosa (L.) Trin. increased linearly (from –0.7 to 11.0) with application rate in the case of U. The response to AN was in the same direction but smaller. While others have shown that the initial response of nitrogen-limited systems to additions of N is a change of ¹⁵N abundance towards that of added N, this study shows that further and excessive additions leads to a retention of ¹⁵N. Monitoring ¹⁵N abundance over time in dose-response trials of this type thus opens new possibilities to estimate critical loads of N and the point of nitrogen saturation.     

Effects of plant breeding and selection on yields and nitrogen fixation in soybeans under two soil nitrogen regimes

Summary Soybeans (Glycine max (L.) Merr.) have a high N requirement which is fulfilled by soil N uptake and N₂-fixation. This study was concerned with the effects of past yield selection on N₂-fixation in soybeans.The soybean cultivars, Lincoln, Shelby, and Williams, which represent successive improvements in the Lincoln germplasm, and a non-nodulating control were planted in a soil containing¹⁵N labelled organic matter. Two replications occurred on soil previously cropped to alfalfa and two on soil previously cropped to soybeans. Plants were harvested at five growth stages and leaf area, plant weight, total N, and atom percent¹⁵N were determined. Mature grain was harvested and yield components were also determined, as well as the total N and¹⁵N content.Cultivar differences in total dry matter were only evident at physiological maturity, when Williams contained the greatest dry matter. Williams exhibited the longest period of seed formation and seed fill and also had the highest grain yield which resulted from a larger weight per seed.The N content of the cultivars did not vary until physiological maturity when Williams contained the highest percent N. The quantity of N fixed at physiological maturity was highest for Williams and lowest for Lincoln. Fixed N contained in the harvested grain was greater for Williams than for the other two cultivars. The fraction of the total plant N derived from fixation was not greatly affected by cultivar and all cultivars acquired an average of 50% of their total N through N₂-fixation.Previous cropping history greatly affected the quantity of N fixed and the fraction of the total plant N derived from fixation. Soybeans following soybeans were more dependent upon N₂-fixation than soybeans following alfalfa with the former deriving 65% of the total plant N from fixation and the latter only 32%. These soybean cultivars apparently utilized soil N first and then used N₂-fixation to satisfy their N requirement.The past selection for higher yield has resulted in soybean cultivars with improved capacities to fix atmospheric N₂ and an improved ability to take up available soil N.     

Evaluation of15N methods to measure nitrogen transfer from alfalfa to companion timothy

Four different methods: direct¹⁵N₂ exposure, legume leaf labeled with¹⁵N,¹⁵N dilution and total N balance were applied to assess the nitrogen transfer (NT) from alfalfa to companion timothy. Evidence of NT was obtained in all cases, which represents about 3% of total N fixed by alfalfa or 10% of N content in timothy at the first cycle of growth. All the three¹⁵N methods gave identical results, while the conventional calculation of NT from the difference of N content in timothy from mixture and monoculture resulted in an over-estimation. The advantages and disadvantages of each method as applied to field conditions are discussed.Contribution No 1158 from the Plant Research Centre.     

Production,standing biomass and natural abundance of <Superscript>15</Superscript>N and <Superscript>13</Superscript>C in ectomycorrhizal mycelia collected at different soil depths in two forest types

Nutrient uptake by forest trees is dependent on ectomycorrhizal (EM) mycelia that grow out into the soil from the mycorrhizal root tips. We estimated the production of EM mycelia in root free samples of pure spruce and mixed spruce-oak stands in southern Sweden as mycelia grown into sand-filled mesh bags placed at three different soil depths (0–10, 10–20 and 20–30 cm). The mesh bags were collected after 12 months and we found that 590±70 kg ha^–1 year^–1 of pure mycelia was produced in spruce stands and 420±160 kg ha^–1 year^–1 in mixed stands. The production of EM mycelia in the mesh bags decreased with soil depth in both stand types but tended to be more concentrated in the top soil in the mixed stands compared to the spruce stands. The fungal biomass was also determined in soil samples taken from different depths by using phospholipid fatty acids as markers for fungal biomass. Subsamples were incubated at 20°C for 5 months and the amount of fungal biomass that degraded during the incubation period was used as an estimate of EM fungal biomass. The EM biomass in the soil profile decreased with soil depth and did not differ significantly between the two stand types. The total EM biomass in the pure spruce stands was estimated to be 4.8±0.9×10³ kg ha^–1 and in the mixed stands 5.8±1.1×10³ kg ha^–1 down to 70 cm depth. The biomass and production estimates of EM mycelia suggest a very long turnover time or that necromass has been included in the biomass estimates. The amount of N present in EM mycelia was estimated to be 121 kg N ha^–1 in spruce stands and 187 kg N ha^–1 in mixed stands. The ¹³C value for mycelia in mesh bags was not influenced by soil depth, indicating that the fungi obtained all their carbon from the tree roots. The ¹³C values in mycelia collected from mixed stands were intermediate to values from pure spruce and pure oak stands suggesting that the EM mycelia received carbon from both spruce and oak trees in the mixed stands. The ¹⁵N value for the EM mycelia and the surrounding soil increased with soil depth suggesting that they obtained their entire N from the surrounding soil.     

Utilization of organic nitrogen at two different substrate pH by different ectomycorrhizal fungi growing in symbiosis with Pinus sylvestris seedlings

The aim of this study was to test the potential of four isolates of ectomycorrhizal (EM) fungi to utilize organic nitrogen (N) at two different substrate pHs. The organic N source (¹⁵N labelled lyophilised fungal mycelium) was mixed with either untreated peat/sand mixture (pH 4.9) or peat/sand mixture limed to a pH of 5.9 and put in cylindrical containers added to each pot. The content of the containers was separated from the roots of Pinus sylvestris seedlings by a nylon mesh and a 2 mm air gap to reduce diffusion of labelled N to the roots. The mycorrhizal plants (except those colonized by Suillus variegatus 2) took up significantly more ¹⁵N from the labelled mycelium than uncolonized seedlings. Liming significantly reduced the uptake of ¹⁵N by one of the EM fungi (unidentified) but not the other tested species (Paxillus involutus and two isolates of S. variegatus). The EM fungal isolates differed in their influence on the bacterial activity of the soil. This was reduced with P. involutus at both pH levels and increased with one of the two S. variegatus isolates at the high pH and with the other S. variegatus isolate at the low pH level. Liming the soil generally increased bacterial activity. The influence of liming on the proportion of organic N uptake in relation to inorganic N uptake by ectomycorrhizal trees is discussed.     

Labeled nitrogen fertilizer research with urea in the semi-arid tropics

Summary As part of a research program to determine the fate of N fertilizers applied to dryland sorghum in the semi-arid tropics,¹⁵N balance studies were conducted with various N sources in the greenhouse. Two American soils, Houston Black clay (Udic Pellustert) and Windthorst sandy loam (Udic Paleustalf), similar in properties to the Vertisol and Alfisol in the semi-arid tropics of India, were employed. Experiments were conducted with large pots containing 20 or 60 kg of soil which was subjected to several watering regimes. The¹⁵N not accounted for in the plant and soil was presumably lost. Losses of N on calcareous Houston Black clay were greatest for broadcast urea, 16%–28%. Amendment of broadcast urea with 2% phenyl phosphorodiamidate, a urease inhibitor, reduced N losses only slightly to 15%–20%. Point placement of urea at a 6 cm soil depth reduced losses to 1%–11%. Granule size had no effect on N loss from point-placed urea. Ammonia volatilization was apparently the main N loss mechanism, since N losses from sodium nitrate were less than 7%, except when the soil surface was waterlogged. N losses on the Windthorst soil averaged 30% from urea and 11% from ammonium nitrate. Amendment of urea with urea phosphate to form a 27% N and 13% P product reduced fertilizer N losses but did not increase grain yield on Windthorst soil. N losses were also less from ammonium nitrophosphate than from urea. Band and point placement of urea 6 cm below the soil surface were equally effective in reducing N loss on Houston Black clay. The findings give credence to the recommendation of deep band placement for urea in the semi-arid tropics.     

Ectomycorrhizal responses to organic and inorganic nitrogen sources when associating with two host species

While it is established that increasing atmospheric inorganic nitrogen (N) deposition reduces ectomycorrhizal fungal biomass and shifts the relative abundances of fungal species, little is known about effects of organic N deposition. The effects of organic and inorganic N deposition on ectomycorrhizal fungi may differ because responses to inorganic N deposition may reflect C-limitation. To compare the effects of organic and inorganic N additions on ectomycorrhizal fungi, and to assess whether host species may influence the response of ectomycorrhizal fungi to N additions, we conducted an N addition experiment at a field site in the New Jersey pine barrens. Seedlings of two host species, Quercus velutina (black oak) and Pinus rigida (pitch pine), were planted at the base of randomly-selected mature pitch pine trees. Nitrogen was added as glutamic acid, ammonium, or nitrate at a rate equivalent to 227.5 kg ha⁻¹ y⁻¹ for eight weeks, to achieve a total application of 35 kg ha⁻¹ during the 10-week study period. Organic and inorganic N additions differed in their effects on total ectomycorrhizal root tip abundance across hosts, and these effects differed for individual morphotypes between oak and pine seedlings. Mycorrhizal root tip abundance across hosts was 90 % higher on seedlings receiving organic N compared to seedlings in the control treatment, while abundances were similar among seedlings receiving the inorganic N treatments and seedlings in the control. On oak, 33–83 % of the most-common morphotypes exhibited increased root tip abundances in response to the three forms of N, relative to the control. On pine, 33–66 % of the most-common morphotypes exhibited decreased root tip abundance in response to inorganic N, while responses to organic N were mixed. Plant chemistry and regression analyses suggested that, on oak seedlings, mycorrhizal colonization increased in response to N limitation. In contrast, pine root and shoot N and C contents did not vary in response to any form of N added, and mycorrhizal root tip abundance was not associated with seedling N or C status, indicating that pine received sufficient N. These results suggest that in situ organic and inorganic N additions differentially affect ectomycorrhizal root tip abundance and that ectomycorrhizal fungal responses to N addition may be mediated by host tree species.     

Carbon and nitrogen mineralization from decomposing gypsy moth frass

Defoliation of forests by insects is often assumed to produce a pulse of available nitrogen (N) from the decomposition of frass pellets. In this study we measured rates of carbon (C) and N mineralization from gypsy moth frass incubated with and without soil, and for soil alone. Incubations were at constant temperature and soil moisture conditions and lasted for 120 days. We found that gypsy moth frass contains much labile C as well as extractable N, and that the stimulation of microbial growth by the labile C results in immobilization of essentially all of the extractable N in the frass. The response of the microbes is fast, beginning within 1 day and lasting at least 90 days. This immobilization response represents an efficient mechanism for conserving N within a forest ecosystem after a defoliation event.     

Mineralization of organically bound nitrogen in soil as influenced by plant growth and fertilization

Summary A loam soil containing an organic fraction labelled with¹⁵N was used for pot experiments with spring barley, rye-grass and clover. The organically bound labelled N was mineralized at a rate corresponding to a half-life of about 9 years. Fertilization with 106 and 424 kgN/ha of unlabelled N in the form of KNO₃ significantly increased uptake of labelled N from the soil in barley and the first harvest of rye-grass crops. The fertilized plants removed all the labelled NH₄ and NO₃ present in the soil, whereas the unfertilized plants removed only about 80%. The second, third and fourth harvests of the unfertilized rye-grass took up more labelled N than the fertilized rye-grass. The total uptake in the four harvests was similar whether the plants were fertilized or not. Application of KCl to barley plants in amounts equivalent to that of KNO₃ resulted in a small but insignificant increase in uptake of labelled N. The uptake of labelled N in the first harvest of clover which was not fertilized but inoculated with Rhizobium was similar to that of the fully fertilized rye-grass indicating that the biological fixation of N had the same effect as addition of N-fertilizer. N uptake in the following harvests was lower and the total uptake by four harvests of clover was similar to that of rye-grass. There was no indication that fertilization with KNO₃ accelerated the mineralization of the organically bound labelled N. The observed apparent ‘priming effect’ of the fertilizer on the uptake of labelled N was compensated by subsequent crops and harvests, and it seems to arise from a more thorough search of the soil volume by a better developed root system of the fertilized plants.     

Carbon and nitrogen utilization and acid production by mycelia of the ectomycorrhizal fungusTricholoma bakamatsutake in vitro

Mycelial growth of an isolate ofT. bakamatsutake was tested in media with C/N ratio ranging from 0 to 50 and with 32 carbon and 12 nitrogen sources. The isolate grew best at the C/N ratio of 30. It utilized the monosaccharidesd-glucose,d-mannose, andd-fructose, the disaccharide trehalose, and polysaccharide pectin among the carbon sources; and yeast extract,l-glutamic acid, and ammonium compounds among the nitrogen sources. The growth of ten isolates and secretion of gluconic and oxalic acids were compared ind-glucose, trehalose, and pectin media. The utilization ofd-glucose, trehalose, and pectin differed among the ten isolates, but all the isolates secreted gluconic acid in thed-glucose media and oxalic acid in the pectin media.     

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.