The vertical distribution of N and K uptake in relation to root distribution and root uptake capacity in mature Quercus robur, Fagus sylvatica and Picea abies stands

We have measured the uptake capacity of nitrogen (N) and potassium (K) from different soil depths by injecting ¹⁵N and caesium (Cs; as an analogue to K) at 5 and 50 cm soil depth and analysing the recovery of these markers in foliage and buds. The study was performed in monocultures of 40-year-old pedunculate oak (Quercus robur), European beech (Fagus sylvatica) and Norway spruce (Picea abies (L.) Karst.) located at an experimental site in Palsgård, Denmark. The markers were injected as a solution through plastic tubes around 20 trees of each species at either 5 or 50 cm soil depth in June 2003. After 65 days foliage and buds were harvested and the concentrations of ¹⁵N and Cs analysed. The recovery of ¹⁵N in the foliage and buds tended to be higher from 5 than 50 cm soil depth in oak whereas they where similar in spruce and beech after compensation for differences in immobilization of ¹⁵N in the soil. In oak more Cs was recovered from 5 than from 50 cm soil depth whereas in beech and spruce no difference could be detected. Out of the three investigated tree species, oak was found to have the lowest capacity to take up Cs at 50 cm soil depth compared to 5 cm soil depth also after compensating for differences in discrimination against Cs by the roots. The uptake capacity from 50 cm soil depth compared with 5 cm was higher than expected from the root distribution except for K in oak, which can probably be explained by a considerable overlap of the uptake zones around the roots and mycorrhizal hyphae in the topsoil. The study also shows that fine roots at different soil depths with different physiological properties can influence the nutrient uptake of trees. Estimates of fine root distribution alone may thus not reflect the nutrient uptake capacity of trees with sufficient accuracy. Our study shows that deep-rooted trees such as oak may have lower nutrient uptake capacity at deeper soil layers than more shallow-rooted trees such as spruce, as we found no evidence that deep-rooted trees obtained proportionally more nutrients from deeper soil layers. This has implications for models of nutrient cycling in forest ecosystems that use the distribution of roots as the sole criterion for predicting uptake of nutrients from different soil depths.     

Do oaks have different strategies for uptake of N, K and P depending on soil depth?

The uptake of nutrients from deep soil layers has been shown to be important for the long-term nutrient sustainability of forest soils. When modelling nutrient uptake in forest ecosystems, the nutrient uptake capacity of trees is usually defined by the root distribution. However, this leads to the assumption that roots at different soil depths have the same capacity to take up nutrients. To investigate if roots located at different soil depths differ in their nutrient uptake capacity, here defined as the nutrient uptake rate under standardized conditions, a bioassay was performed on excised roots (<1 mm) of eight oak trees (Quercus robur L.). The results showed that the root uptake rate of ⁸⁶Rb⁺ (used as an analogue for K⁺) declined with increasing soil depth, and the same trend was found for . The root uptake rate of , on the other hand, did not decrease with soil depth. These different physiological responses in relation to soil depth indicate differences in the oak roots, and suggest that fine roots in shallow soil layers may be specialized in taking up nutrients such as K⁺ and which have a high availability in these layers, while oak roots in deep soil layers are specialized in taking up other resources, such as P, which may have a high availability in deep soil layers. Regardless of the cause of the difference in uptake trends for the various nutrients, these differences have consequences for the modelling of the soil nutrient pool beneath oak trees and raise the question of whether roots can be treated uniformly, as has previously been done in forest ecosystem models. Responsible Editor: Herbert Johannes Kronzucker.     

The Vertical Pattern of Rooting and Nutrient Uptake at Different Altitudes of a South Ecuadorian Montane Forest

The vertical pattern of root length densities (RLD) of fine roots (<2 mm in diameter) and nitrogen (N) uptake potential were determined at different altitudes (1,900, 2,400, and 3,000 m a.s.l.) of a tropical montane forest in order to improve our knowledge about the depth distribution of nutrient uptake in this ecosystem. At higher altitudes, precipitation rate and frequency of fog were higher than at lower altitudes while mean annual air temperature decreased with increasing altitude. Soils were always very acid with significantly lower pH at a depth of 0.0–0.3 m in mineral soil at 3,000 m (2.8–2.9) than at 1,900 and 2,400 m (3.1–3.5). The vertical distribution of RLD was very similar both during the dry and the rainy season. During the dry season the percentage of root length in the organic layer increased from 51% at 1,900 m to 61% at 2,400 m and 76% at 3,000 m. At 3,000 m, RLD was markedly higher in the upper 0.05 m than in the remaining organic layer, whereas at 1,900 m and 2,400 m RLD were similar in all depths of the organic layer. In mineral soil, RLD decreased to a greater degree with increasing soil depth at the upper two study sites than at 1,900 m. The relative N uptake potential from different soil layers (RNUP) was determined by ¹⁵N enrichment of leaves after application of ¹⁵N enriched ammonium sulphate at various soil depths. RNUP closely followed fine root distribution confirming the shallower pattern of nutrient uptake at higher altitudes. RNUP was very similar for trees, shrubs and herbs, but shallower for saplings which obtained N only from the organic layer at both altitudes. Liming and fertilizing (N, P, K, Mg) of small patches in mineral soil had no significant impact on fine root growth. We conclude that the more superficial nutrient uptake ability at higher altitudes may be partly related to increased nutrient input from canopy by leaching. However, the specific constraints for root growth in the mineral soil of tropical montane forests warrant further investigations.     

Root competition for phosphorus between the tree and herbaceous components of silvopastoral systems in Kerala, India

Root competition in polyculture systems involving combinations of four tree species and four grass species was evaluated based on ³²P recovery by each species in mixed and sole crop situations. The tree species were: Leucaena leucocephala, Casuarina equisetifolia, Acacia auriculiformis and Ailanthus triphysa, and the grass species were: Pennisetum purpureum (hybrid napier), Brachiaria ruziziensis (congo signal), Panicum maximum (guinea grass) and Zea mexicana (teosinte). Four lateral distance (25 and 50 cm) and depth (15 and 50 cm) treatments were included in the study to characterize the relative fine root distribution of trees. Absorption of ³²P was monitored through radioassay of leaves. Regardless of the species, ³²P uptake from 50 cm soil depth was lower than that of 15 cm depth. Absorption of ³²P from 50 cm lateral distance was also less than that of 25 cm distance in Acacia and Casuarina. Grass species in sole crop situations absorbed more ³²P than in mixed systems. None of the grass species when grown in association with tree components affected the absorption of ³²P by trees. All grass species exerted a complementary effect on ³²P absorption by Casuarina. Leucaena also benefited in the same way when grown in association with congo signal and/or teosinte. Of the tree species, Acacia and Leucaena adversely affected the ³²P uptake by grass species.     

Estimating the relative nutrient uptake from different soil depths in Quercus robur, Fagus sylvatica and Picea abies

The distribution of fine roots and external ectomycorrhizal mycelium of three species of trees was determined down to a soil depth of 55 cm to estimate the relative nutrient uptake capacity of the trees from different soil layers. In addition, a root bioassay was performed to estimate the nutrient uptake capacity of Rb⁺ and NH₄⁺ by these fine roots under standardized conditions in the laboratory. The study was performed in monocultures of oak (Quercus robur L.), European beech (Fagus sylvatica L.) and Norway spruce [Picea abies (L.) Karst.] on sandy soil in a tree species trial in Denmark. The distribution of spruce roots was found to be more concentrated to the top layer (0–11 cm) than that of oak and beech roots, and the amount of external ectomycorrhizal mycelia was correlated to the distribution of the roots. The uptake rate of [⁸⁶Rb⁺] by oak roots declined with soil depth, while that of beech or spruce roots was not influenced by soil depth. In modelling the nutrient sustainability of forest soils, the utilization of nutrient resources in deep soil layers has been found to be a key factor. The present study shows that the more shallow-rooted spruce can have a similar capacity to take up nutrients from deeper soil layers than the more deeply rooted oak. The distribution of roots and mycelia may therefore not be a reliable parameter for describing nutrient uptake capacity by tree roots at different soil depths.     

Fertilizer and soil nitrogen accumulation by irrigated barley grown on a red-brown earth in south-eastern Australia

¹⁵N labelled (NH₄)₂SO₄ was applied to barley at 5 g N m⁻² (50 kg N ha⁻¹) in microplots at sowing to study the timing of the N losses and the contribution of soil and fertilizer N to the plant. Water treatments included rainfed and irrigation at 45–50 mm deficit beginning in the spring. Recovery of¹⁵N in the plant increased to a maximum of about 20% within 91 days after sowing (DAS 91) and then remained constant. Approximately 16% (0.8 g N m⁻²) of the fertilizer was in the stem and leaves at DAS 91 and this N was subsequently redistributed to the head. At maturity, approximately 75% of the¹⁵N assimilated by the tops was recovered in the grain. Soil N contributed 3.6 g N m⁻² to the head; 2.2 g N m⁻² was remobilized from the stem and leaves, and the balance, approximately 1.4 g N m⁻², was taken up from the soil between DAS 69 to 91. Effects of irrigation treatments on N accumulation were not significant. Residual¹⁵N fertilizer in the soil decreased with time from sowing, and at maturity 40% of the applied N was recovered in the surface 0.15 m.¹⁵N movement to depth was limited and less than 5% of the fertilizer was recovered below 0.15 m. Irrigation had no effect on the¹⁵N recovery at depth. Total recovery of the¹⁵N varied between 60 and 67% and implies that 33–40% was lost from the soil-plant system. The total recovery in the soil and plant was not affected by time or irrigation in the interval DAS 39 to 134. Losses occurred before DAS 39 when crop uptake of N was small and soil mineral N content was high. There was an apparent loss of 1.9 g fertilizer N m⁻² (i.e. 38% of that applied) between DAS 1 and 15. This loss occurred before crop emergence when rainfall provided conditions suitable for denitrification.     

Temporal and spatial distribution of roots and competition for nitrogen in pea-barley intercrops – a field study employing 32P technique

Root system dynamics, productivity and N use were studied in inter- and sole crops of field pea (Pisum sativum L.) and spring barley (Hordeum vulgare L.) on a temperate sandy loam. A ³²P tracer placed at a depth of 12.5, 37.5, 62.5 or 87.5 cm was employed to determine root system dynamics by sampling crop leaves at 0, 15, 30 and 45 cm lateral distance. ¹⁵N addition was used to estimate N₂ fixation by pea, using sole cropped barley as reference crop. The Land Equivalent Ratio (LER), which is defined as the relative land area under sole crops that is required to produce the yields achieved in intercropping, were used to compare the crop growth in intercrops relative to the respective sole crops.The ³²P appearance in leaves revealed that the barley root system grows faster than that of pea. P uptake by the barley root system during early growth stages was approximately 10 days ahead of that of the pea root system in root depth and lateral root distribution. More than 90% of the P uptake by the pea root system was confined to the top 12.5 cm of soil, whereas barley had about 25–30% of tracer P uptake in the 12.5 – 62.5 cm soil layer. Judging from this P uptake, intercropping caused the barley root system to grow deeper and faster lateral root development of both species was observed. Barley accumulated similar amounts of aboveground N when grown as inter- and sole crop, whereas the total aboveground N acquired by pea in the intercrop was only 16% of that acquired in the pea sole crop. The percentage of total aboveground N derived from N₂ fixation in sole cropped pea increased from 40% to 80% during the growth period, whereas it was almost constant at 85% in intercropped pea. The total amounts of N₂ fixed were 95 and 15 kg N ha^–1 in sole cropped and intercropped pea, respectively. Barley was the dominant component of the pea-barley intercrop, obtaining 90% of its sole crop yield, while pea produced only 15% of the grains of a sole crop pea. Intercropping of pea and barley improved the utilization of plant growth resources (LER > 1) as compared to sole crops. Root system distribution in time and space can partly explain interspecific competition. The ³²P methodology proved to be a valuable tool for determining root dynamics in intercropping systems.     

Some observations on the root growth of young apple trees and their uptake of nutrients when grown in herbicided strips in grassed orchards

Summary Root laboratory observations of the root growth of 4-year-old trees of Cox/M.26 planted in a herbicided strip in grass indicated that during the year 70% of the new growth occurred in the strip. Growth appeared to begin earlier during the year under the bare soil than under grass. Nitrogen absorption from the strip and the grassed alley was assessed by measuring ¹⁵N uptake; at 10 cm depth uptake was almost entirely from the strip. An experiment using 2-year-old trees of Cox/MM. 106 and ¹⁵N placements at 7.5 and 15 cm depths in the strip and 15 cm depth in the grassed alley gave similar results. With ³²P as a tracer and similar trees a small amount of uptake from 25 cm depth under grass was detected. The experiments indicate that young trees produce most of their new roots in the herbicide strips where most of their nutrient uptake occurs and little or none from the grassed alleys. The absorption of nitrogen into the leaves was greater in early summer than autumn. re]19751017     

Regulation of nitrogen partitioning in field-grown almond trees: Effects of fruit load and foliar nitrogen applications

Two treatments were employed to influence the amount of amino nitrogen (N) transport in phloem. In walnut trees (Juglans regia L.), developing fruit significantly reduced the efflux of foliar-applied ¹⁵N-enriched urea from treated spurs over a 33-day period in comparison with similarly-treated defruited spurs. Those data suggest that local aboveground demand for N influences vascular transport of amino N. In another experiment, a 1% urea solution was applied foliarly to 5-year old `Mission' almond trees [Prunus dulcis (Mill.) D. A. Webb] to increase the concentration of amino N in the phloem. The effect of foliar N treatments on a) the transport and distribution of labelled urea N within the trees over the experimental period and b) the uptake of soil-applied labelled N were determined by replicated whole tree excavation, fractionation into various tree components and mass spectrometric analyses of the ¹⁴N/¹⁵N ratios. Concentrations and composition of amino acids in the phloem and xylem saps of control trees and trees receiving foliar-applied urea were also determined. In foliar urea-treated trees, the amino acid concentrations increased significantly in leaf and bark phloem exudate, within 24 and 96 h, respectively. Foliar-applied urea N was translocated to the roots of almond trees over the experimental period and decreased soil N uptake. The results of these experiments are consistent with the hypothesis that aboveground N demand affects the amount of amino N cycling between shoots and roots, and may be involved in the regulation of soil N uptake. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

Root distributions partially explain 15N uptake patterns in Gliricidia and Peltophorum hedgerow intercropping systems

The relative distributions of tree and crop roots in agroforestry associations may affect the degree of complementarity which can be achieved in their capture of below ground resources. Trees which root more deeply than crops may intercept leaching nitrogen and thus improve nitrogen use efficiency. This hypothesis was tested by injection of small doses of (¹⁵NH₄)₂SO₄ at 21.8 atom% ¹⁵N at different soil depths within established hedgerow intercropping systems on an Ultisol in Lampung, Indonesia. In the top 10 cm of soil in intercrops of maize and trees, root length density (L_rv) of maize was greater than that of Gliricidia sepium trees, which had greater L_rv in this topsoil layer than Peltophorum dasyrrachis trees. Peltophorum trees had a greater proportion of their roots in deeper soil layers than Gliricidia or maize. These vertical root distributions were related to the pattern of recovery of ¹⁵N placed at different soil depths; more ¹⁵N was recovered by maize and Gliricidia from placements at 5 cm depth than from placements at 45 or 65 cm depth. Peltophorum recovered similar amounts of ¹⁵N from placements at each of these depths, and hence had a deeper N uptake distribution than Gliricidiaor maize. Differences in tree L_rv across the cropping alley were comparatively small, and there was no significant difference (P<0.05) in the uptake of ¹⁵N placed in topsoil at different distances from hedgerows. A greater proportion of the ¹⁵N recovered by maize was found in grain following ¹⁵N placement at 45 cm or 65 cm depth than following placement at 5 cm depth, reflecting the later arrival of maize roots in these deeper soil layers. Thus trees have an important role in preventing N leaching from subsoil during early crop establishment, although they themselves showed a lag phase in ¹⁵N uptake after pruning. Residual ¹⁵N enrichment in soil was strongly related to application depth even 406 days after ¹⁵N placement, demonstrating the validity of this approach to mapping root activity distributions.     

Authors index

In a runoff irrigation system in Northern Kenya, we studied the nutrient interactions of sole cropped and alley cropped Sorghum bicolor (L.) Moench and Acacia saligna (Labill.) H.L. Wendl. The trees were pruned once before the cropping season and the biomass was used as fodder for animals. The nutrient contents in leaf tissue, soil and soil solution were monitored and the uptake of applied tracers (¹⁵N, Sr) was followed. The grain yield of alley cropped sorghum was similar to or slightly higher than in monoculture and did not decrease near the tree-crop interface. Foliar N and Ca contents of the crop were higher in the agroforestry combination than in monoculture, corresponding to higher soil N and Ca contents. Soil solution and soil mineral N dynamics indicate an increase of N under the tree row and unused soil N at the topsoil in the alley of the sole cropped trees as well as below 60 cm depth in the crop monoculture. The N use efficiency of the tree+crop combination was higher than the sole cropped trees or crops. Competition was observed for Zn and Mn of both tree and crop whereas for Ca only the tree contents decreased. P, K, Mg and Fe dynamics were not affected by alley cropping at our site. The lower uptake of applied Sr by trees in alley cropping compared to those of the monoculture stand suggested a lower competitiveness of the acacia than sorghum, which did not show lower Sr contents when intercropped. The study showed the usefulness of combining soil and plant analyses together with tracer techniques identifying nutrient competition, nutrient transfer processes and the complementary use of soil nutrients, as the main features of the tree-crop combination. This revised version was published online in June 2006 with corrections to the Cover Date.     

A bioassay of the availability of residual 15N fertilizer eight years after application to a forest soil in interior British Columbia

Although a high proportion of fertilizer N may be immobilized in organic forms in the soil, no studies have examined the long-term availability of residual fertilizer ¹⁵N in forestry situations. We investigated this by growing lodgepole pine (Pinus contorta) seedlings in surface (0–10 cm) soil sample eight years after application of ¹⁵N-urea, ¹⁵NH₄NO₃ and NH₄ ¹⁵NO₃ to lodgepole pine in interior British Columbia. After nine months of growth in the greenhouse, seedlings took up an average of 8.5% of the ¹⁵N and 4.6% of the native N per pot. Most of the mineral N in the pots without seedlings was in the form of nitrate, while pots with seedlings had very low levels of mineral N. In contrast to the greenhouse study, there was no significantuptake of ¹⁵N by trees in the field study after the first growing season, although half of the soil organic ¹⁵N was lost between one and eight years after fertilization. This indicates the need to understand the mechanisms which limit the uptake of mineral N by trees in the field, and the possible mismatch of tree demand and mineral N availability.     

Transformations and movement of P in the rhizosphere

Summary Wheat plants labelled with³³P were grown in thin layers of soil amended with³²P-labelled fertiliser. Roots were separated from the soil during plant growth by a porous membrane to overcome difficulties in measuring microbial P in rhizosphere soil. Over the 22 day growth period, net movement of³³P out of healthy growing roots varied from 0.9–4.9% of the total³³P translocated to the root. Over the same period the plants took up 12.0% and the microbial biomass 14.1% of the fertiliser³²P. On drying and rewetting of the soil after the plants were harvested, a large proportion of root P moved into soil fractions while³²P appeared to accumulate in the biomass and stable P forms.     

Estimation of biological nitrogen fixation associated withBrachiaria andPaspalum grasses using15N labelled organic matter and fertilizer

Summary Six pasture grasses,Paspalum notatum cv batatais,P. notatum cv pensacola,Brachiaria radicans, B. ruziziensis, B. decumbens andB. humidicola, were grown in concrete cylinders (60 cm diameter) in the field for 31 months. The soil was amended with either a single addition of¹⁵N labelled organic matter or frequent small (2 kg N. ha^–1) additions of¹⁵N enriched (NH₄)₂SO₄. In the labelled fertilizer treatment soil analysis revealed that there was a very drastic change in¹⁵N enrichment in plant-available nitrogen (NO ₃ ^– +NH ₄ ⁺ ) with depth. The different grass cultivars recovered different quantities of applied labelled N, and evidence was obtained to suggest that the roots exploited the soil to different depths thus obtaining different¹⁵N enrichments in soil derived N. This invalidated the application of the isotope dilution technique to estimate the contribution of nitrogen fixation to the grass cultivars in this treatment. In the labelled organic matter treatment the¹⁵N label in the plant-available N declined at a decreasing rate during the experiment until in the last 12 months the decrease was only from 0.274 to 0.222 atom % excess. There was little change in¹⁵N enrichment of available N with depth, hence it was concluded that although the grasses recovered different quantities of labelled N, they all obtained virtually the same¹⁵N enrichment in soil derived N. Data from the final harvests of this treatment indicated thatB. humidicola andB. decumbens obtained 30 and 40% respectively of their nitrogen from N₂ fixation amounting to an input of 30 and 45 kg N.ha^–1 year^–1 respectively.     

Carbon, nitrogen and phosphorus in volcanic soils following afforestation with native birch (Betula pubescens) and introduced larch (Larix sibirica) in Iceland

Afforestation has become an important tool for soil protection and land reclamation in Iceland. Nevertheless, the harsh climate and degraded soils are growth-limiting for trees, and little is know about changes in soil nutrients in maturing forests planted on the volcanic soils. In the present chronosequence study, changes in C, N and total P in soil (0–10 and 10–20 cm depth) and C and N in foliar tissue were investigated in stands of native Downy birch (Betula pubescens Enrh.) and the in Iceland introduced Siberian larch (Larix sibirica Ledeb.). The forest stands were between 14 and 97 years old and were established on heath land that had been treeless for centuries. Soils were Andosols derived from basaltic material and rhyolitic volcanic ash. A significant effect of tree species was only found for the N content in foliar tissue. Foliar N concentrations were significantly higher and foliar C/N ratios significantly lower in larch needles than in birch leaves. There was no effect of stand age. Changes in soil C and the soil nutrient status with time after afforestation were little significant. Soil C concentrations in 0–10 cm depth in forest stands older than 30 years were significantly higher than in heath land and forest stands younger than 30 years. This was attributed to a slow accumulation of organic matter. Soil N concentrations and soil P_tot were not affected by stand age. Nutrient pools in the two soil layers were calculated for an average weight of soil material (400 Mg soil ha⁻¹ in 0–10 cm depth and 600 Mg soil ha⁻¹ in 10–20 cm depth, respectively). Soil nutrient pools did not change significantly with time. Soil C pools were in average 23.6 Mg ha⁻¹ in the upper soil layer and 16.9 Mg ha⁻¹ in the lower soil layer. The highest annual increase in soil C under forest compared to heath land was 0.23 Mg C ha⁻¹ year⁻¹ in 0–10 cm depth calculated for the 53-year-old larch stand. Soil N pools were in average 1.0 Mg N ha⁻¹ in both soil layers and did not decrease with time despite a low N deposition and the uptake and accumulation of N in biomass of the growing trees. Soil P_tot pools were in average 220 and 320 kg P ha⁻¹ in the upper and lower soil layer, respectively. It was assumed that mycorrhizal fungi present in the stands had an influence on the availability of N and P to the trees. Responsible Editor: Hans Lambers.     

Competition in tree row agroforestry systems. 2. Distribution, dynamics and uptake of soil inorganic N

The effect of tree row species on the distribution of soil inorganic N and the biomass growth and N uptake of trees and crops was investigated beneath a Grevillea robustaA. Cunn. ex R. Br. (grevillea) tree row and Senna spectabilisDC. (senna) hedgerow grown with Zea mays L. (maize) and a sole maize crop, during one cropping season. The hypothesis was that a tree with a large nutrient uptake would have a greater competitive effect upon coexisting plants than a tree that takes up less and internally cycles nutrients. The field study was conducted on a kaolinitic Oxisol in the sub-humid highlands of western Kenya. Soil nitrate and ammonium were measured to 300 cm depth and 525 cm distance from the tree rows, before and after maize cropping. Ammonium concentrations were small and did not change significantly during the cropping season. There was > 8 mg nitrate kg^–1 in the upper 60 cm and at 90–180 cm depth at the start of the season, except within 300 cm of the senna hedgerow where concentrations were smaller. During the season, nitrate in the grevillea-maize system only decreased in the upper 60 cm, whereas nitrate decreased at almost every depth and distance from the senna hedgerow. Inorganic N (nitrate plus ammonium) decreased by 94 kg ha^–1 in the senna-maize system and 33 kg ha^–1 in the grevillea-maize system.The aboveground N content of the trees increased by 23 kg ha^–1 for grevillea and 39 kg ha^–1 for senna. Nitrogen uptake by maize was 85 kg ha^–1 when grown with grevillea and 65 kg ha^–1 with senna. Assuming a mineralisation input of 50 kg N ha^–1season^–1, the decrease in inorganic soil N approximately equalled plant N uptake in the grevillea-maize system, but exceeded that in the senna-maize system. Pruning and litter fall removed about 14 kg N ha^–1 a^–1 from grevillea, and > 75 kg N ha^–1 a^–1 from senna. The removal of pruned material from an agroforestry system may lead to nutrient mining and a decline in productivity.     

Native soil and fresh fertilizer phosphorus uptake as affected by rate of application and P fertilizers

³²P labelled fertilizers were used to measure native soil and fresh fertilizer phosphorus uptake byLolium perenne L. in greenhouse experiments. The P source evaluation was carried out for multiple rates of application for a standard P fertilizer (DAP) on low and medium soil P levels and for North Carolina rock phosphate (RP) at the medium soil P level only. At the low soil P level, the native P uptake increase was independent of P-DAP applied, and represented 19% of the nil P uptake. At the medium soil P level, the variability of the native soil as a nutrient P source depended on the phosphate fertilizer applied, and the rate of application. Consequently the amount of total P uptake could conceal differences in P fertilizer evaluations as the nutrient P source. Fresh P uptake increased linearly with the rates of P applied as standard or tested P fertilizer. The comparison of various P sources by means of fresh P uptake ratio (i.e. fresh P uptake from tested phosphate divided by fresh P uptake from standard phosphate) was independent of the rate of application. It was therefore suggested that various phosphorus sources be evaluated by measuring the fresh P uptake for a single rate of application.     

Mobilization of the reserve N in citrus

The mobilization of N from reserve organs (leaves, roots, branches and trunk) to developing new organs was studied at different moments of the growth cycle. Three-year-old Valencia Late orange trees (Citrus sinensis (L.) Osbeck) were grown individually outdoors in 150 L containers filled with siliceous sand. Trees were irrigated with a nutrient solution labelled with potassium nitrate with an enrichment of 4 atom ¹⁵N % excess during a complete growth cycle. At the following year, plants were irrigated with unlabelled nutrient solution, and harvested throughout the growth cycle (flowering, fruit set, second flush, third flush, and dormancy). Total N and ¹⁵N analyses were carried out in the different organs of the plants.The highest amounts of N were found in leaves and roots (33–42% and 30–38%, respectively). Distribution of ¹⁵N was similar to that obtained for total N (42 and 39% of total ¹⁵N in leaves and roots, respectively) as is expected after a long period of labelling. Old leaves were the main reserve organs, contributing a 40–50% of the total N exported. Roots and aerial woody tissues exported between 30–35% and 15–25% of total reserve N, respectively. N exported by old leaves was approximately 57% of the N accumulated during the preceding year, while roots translocated 40% and trunk plus branches 35%. More than 70% of N accumulated in new organs during spring came from N stored in old organs.     

Capture and allocation of nitrogen byQuercus douglasii seedlings in competition with annual and perennial grasses

Summary The spatial overlap of woody plant root systems and that of annual or perennial grasses promotes competition for soil-derived resources. In this study we examined competition for soil nitrogen between blue oak seedlings and either the annual grassBromus mollis or the perennial grassStipa pulchra under controlled outdoor conditions. Short-term nitrogen competition was quantified by injecting¹⁵N at 30 cm depth in a plane horizontal to oak seedling roots and that of their neighbors, and calculating¹⁵N uptake rates, pool sizes and¹⁵N allocation patterns 24 h after labelling. Simultaneously, integrative nitrogen competition was quantified by examining total nitrogen capture, total nitrogen pools and total nitrogen allocation.Stipa neighbors reduced inorganic soil nitrogen content to a greater extent than didBromus plants. Blue oak seedlings responded to lower soil nitrogen content by allocating lower amounts of nitrogen per unit of biomass producing higher root length densities and reducing the nitrogen content of root tissue. In addition, blue oak seedlings growing with the perennial grass exhibited greater rates of¹⁵N uptake, on a root mass basis, compensating for higher soil nitrogen competition inStipa neighborhoods. Our findings suggest that while oak seedlings have lower rates of nitrogen capture than herbaceous neighbors, oak seedlings exhibit significant changes in nitrogen allocation and nitrogen uptake rates which may offset the competitive effect annual or perennial grasses have on soil nitrogen content.