The effects of water content of the topsoil on micronutrient availability and uptake in a siliceous sandy soil

Summary Effects of water content of the topsoil on the supply of manganese, zinc and copper to plants from an acid, siliceous sandy soil were investigated. The experiment was designed to provide a good supply of water and of nutrients other than those mentioned above at all times.Water content of the topsoil affected the root growth in that layer significantly. However, drying of the topsoil for as long as 42 days did not significantly affect the concentration of micronutrients in the shoots of oats, provided the plants did not experience water stress.Despite the acidic, siliceous sandy nature of the topsoil it showed a surprisingly high ability to render added manganese, zinc and copper unavailable. This phenomenon, probably mediated by organic matter, had a dominant effect on the exchangeable micronutrients in comparison to the effects due to fluctuations in soil water content.     

Stable isotope analysis of water extraction from subsoil in upland rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.) as affected by drought and soil compaction

Water extraction from subsoil in upland rice (Oryza sativa L.) was examined as related to topsoil desiccation and subsoil compaction. The water extraction was observed by measurements of heavy water concentrations in transpiring plants. The plants were grown in pots that were filled with sandy soil and vertically compartmented into two columns. Heavy water was applied to the subsoil. Plants exposed to mild topsoil desiccation (–120 kPa in water potential) eventually increased water extraction from the subsoil and maintained photosynthetic rate and stomatal conductance at the wet condition level. The rates of the plants subjected to severely droughted topsoil (–190 kPa) were significantly lowered due to less water uptake from the subsoil. Subsoil compaction at bulk densities of 1.45 and 1.50 Mg m^–3 inhibited increase of root length densities. Limited water extraction from the subsoil was insufficient to maintain plant productivity under drought conditions. Daily water uptake per unit of root length in the lower tube did not apparently increase even if water demand on the unit root length increased. When water to topsoil was completely withheld, water extraction from the subsoil gradually increased as the topsoil dried out. Plants that were watered and rewatered took up very little water from the subsoil. The extraction from the subsoil occurred only when water potential of the topsoil was below about –190 kPa.     

Zinc,Iron, Manganese and Copper Uptake Requirement in Response to Nitrogen Supply and the Increased Grain Yield of Summer Maize

The relationships between grain yields and whole-plant accumulation of micronutrients such as zinc (Zn), iron (Fe), manganese (Mn) and copper (Cu) in maize (Zea mays L.) were investigated by studying their reciprocal internal efficiencies (RIEs, g of micronutrient requirement in plant dry matter per Mg of grain). Field experiments were conducted from 2008 to 2011 in North China to evaluate RIEs and shoot micronutrient accumulation dynamics during different growth stages under different yield and nitrogen (N) levels. Fe, Mn and Cu RIEs (average 64.4, 18.1and 5.3 g, respectively) were less affected by the yield and N levels. ZnRIE increased by 15% with an increased N supply but decreased from 36.3 to 18.0 g with increasing yield. The effect of cultivars on ZnRIE was similar to that of yield ranges. The substantial decrease in ZnRIE may be attributed to an increased Zn harvest index (from 41% to 60%) and decreased Zn concentrations in straw (a 56% decrease) and grain (decreased from 16.9 to 12.2 mg kg⁻¹) rather than greater shoot Zn accumulation. Shoot Fe, Mn and Cu accumulation at maturity tended to increase but the proportions of pre-silking shoot Fe, Cu and Zn accumulation consistently decreased (from 95% to 59%, 90% to 71% and 91% to 66%, respectively). The decrease indicated the high reproductive-stage demands for Fe, Zn and Cu with the increasing yields. Optimized N supply achieved the highest yield and tended to increase grain concentrations of micronutrients compared to no or lower N supply. Excessive N supply did not result in any increases in yield or micronutrient nutrition for shoot or grain. These results indicate that optimized N management may be an economical method of improving micronutrient concentrations in maize grain with higher grain yield.     

Does hydraulic lift exist in shallow-rooted species? A quantitative examination with a half-shrub Gutierrezia sarothrae

Hydraulic lift occurs in some deep-rooted shrub and herbaceous species. In this process, water taken up by deep roots from the moist subsoil is delivered to the drier topsoil where it is later reabsorbed by shallow roots. However, little is known about the existence of hydraulic lift in shallow-rooted xeric species. The objectives of this study were 1) to ascertain whether hydraulic lift exists in Gutierrezia sarothrae (broom snakeweed), a widespread North American desert species with a shallow root system, grown in pot and field conditions and 2) if it does, how much water can be transferred from the subsoil to the 30 cm topsoil during the night. Snakeweed seedlings were transplanted in buried pots allowing the deeper roots to grow into the subsoil 30 cm below the surface. Soil water content inside and outside of the pot was measured seasonally and diurnally with time domain reflectometry technique (TDR). An increase in water content was detected in the pot after the plant was covered for 3 h by an opaque plastic bag during the day, suggesting hydraulic lift from deeper depths and exudation of water into the drier topsoil. Root exudation was also observed on native range sites dominated by snakeweed. Water efflux in the pot was 271 g per plant per night. which was equivalent to 15.3% of the extrapolated, porometer-derived whole-plant daily transpiration. Hydraulic lift observed in Gutierrezia improved water uptake during the day when evaporative demand is high and less water is available in the topsoil. We concluded that hydraulic lift might help snakeweed to alleviate the effect of water stress.     

Tolerance to acid soil conditions of the velvet beans Mucuna pruriens var. utilis and M. deeringiana

Fast growing, climbing leguminous cover crops such as the velvet beans can be used to reclaim weed-infested, degraded soils in the humid tropics, especially land covered by the grass Imperata cylindrica; they climb over the grass leaves and shade the grass out if their cover lasts long enough. Tolerance of two species of velvet bean to eroded soils was investigated by removing topsoil and directly sowing into the subsoil; plots where topsoil was not removed were used as a control. The response to small amounts of P fertilizer and lime was also tested. Removal of the topsoil resulted in retarded growth of both species, in increased dry matter content of the shoot, in decreased specific leaf area and in increased leaf weight ratio, due to shorter internodes. Six weeks after planting the leaf area index (LAI) was about 1.2 where topsoil was retained, sufficient for a shading effect on Imperata. Where topsoil had been removed, the LAI was only 0.6. Mucuna pruriens var. utilis showed a faster aboveground growth than M. deeringiana; the species did not differ in tolerance to eroded soil. Small amounts of P fertilizer had no significant effect on the growth of both Mucuna species. Shoot: root ratios, on a dry weight basis, were much lower when topsoil had been removed, about 3.7 and 2.4 for M. p. utilis and M. deeringiana respectively, compared to 6.2 and 3.3 where topsoil was retained. Removal of topsoil led to reduced Mg and to increased Al concentrations in roots, and to increased levels of Mn and Al in shoots. In the second year no effect of lime or residual effect of P application was found on growth of Mucuna or Imperata. Removal of the topsoil had little effect on the growth of weeds after the cover crop had been harvested. Due to the high Al tolerance of Imperata, reclamation by Mucuna will be less effective if the topsoil has been lost by erosion.     

Some effects of micronutrients and liming on the yield,quality and micronutrient status of tomatoes grown in peat

Summary The effects of a fritted micronutrient mixture on the growth and yield of tomatoes were studied in relation to the pH of the peat substrate. The treatments consisted of three rates of application of the fritted micronutrient mixture (0, 200, 500 g/m³ F 253A) in factorial combination with four rates of liming (average pH values 4.5, 5.1, 5.8 and 6.7). Three crops were grown to maturity, and in each case the total yield of fruit decreased (18–24%) when the peat was limed to pH>6.0 (P<0.001). Omission of the micronutrient mixture reduced the yield by 20–65% (P<0.001). The combination of heavy liming and omission of micronutrients was particularly detrimental, and depressed the yield by 40–87% as compared with the most favourable treatment. The incidence of ripening disorders was also increased by heavy liming and by omission of micronutrients. The boron, copper, iron, manganese and zinc contents of the leaves are tabulated. Multiple regression analyses suggested that the yield was consistently depressed by deficiencies of boron and copper, and responses to iron and manganese were also found.     

Hydraulic properties of layered soils influence survival of Rhodes grass (<Emphasis Type="Italic">Chloris gayana</Emphasis> Kunth.) during water stress

Survival of vegetation on soil-capped mining wastes is often impaired during dry seasons due to the limited amount of water stored in the shallow soil capping. Growth and survival of Rhodes grass (Chloris gayana) during soil drying on various layered capping sequences constructed of combinations of topsoil, subsoil, seawater-neutralised residue sand and low grade bauxite was determined in a glasshouse. The aim was to describe the survival of Rhodes grass in terms of plant and soil water relationships. The soil water characteristic curve and soil texture analysis was a good predictor of plant survival. The combination of soil with a high water holding capacity and low soil water diffusivity (e.g. subsoil with high clay contents) with soil having a high water holding capacity and high diffusivity (e.g. residue sand) gave best survival during drying down (up to 88 days without water), whereas topsoil and low grade bauxite were unsuitable (plants died within 18–39 days). Clayey soil improved plant survival by triggering a water stress response during peak evaporative water demand once residue sand dried down and its diffusivity fell below a critical range. Thus, for revegetation in seasonally dry climates, soil capping should combine one soil with low diffusivity and one or more soils with high total water holding capacity and high diffusivity.     

Potential importance of the subsoil for the P and Mg nutrition of wheat

A method is described which allowed the quantification of the potential uptake of P and Mg from the subsoil (>30cm) by spring wheat. Wheat was grown on an artificial topsoil (sand with no plant available P or Mg) which was superimposed on loess subsoils in N. Germany. The supply of P and Mg in the topsoil was varied by application of different quantities of P and Mg fertilizer. Uptake of P and Mg from the subsoil was calculated as the difference between total plant uptake (determined by plant analysis) and the quantities of P and Mg removed from the topsoil (determined by soil analysis). P uptake from the subsoil increased from 37% to 85% of total P uptake, with decreasing P supply in the topsoil. Calculations of potential supply by diffusion showed that, with a CAL-extractable P₂O₅ content in the subsoil of 9 mg 100g^-1, supply from the subsoil was only possible if the influence of root hairs was considered. The method also showed that the total demand for Mg by spring wheat could be satisfield from the supply of Mg from the subsoil of typical loess soils. Mg uptake from the subsoil decreased to 33% of total uptake with increasing Mg supply in the topsoil.     

Alfalfa,Medicago sativa L., in highly weathered,acid soils

Summary Rooting into acid subsoils would be a desirable trait for alfalfa which should result in better water extraction and yield. In this study, the rooting depth into acid subsoils and top yield of alfalfa plants selected for acid tolerance were investigated in a repacked profile with a limed, fertile topsoil, but unamended, acid subsoil. The effects of subsoil modification by CaCO₃ and CaSO₄·2H₂O addition on alfalfa rooting, top growth, and water extraction were also studied. Plants from acid selections rooted deeper into acid subsoil when compared to control plants (selected under limed conditions). However, the reverse response was found in the CaSO₄·2H₂O treated subsoil. There were no differences among selections for total top yield for any subsoil treatment. Water extraction from the lower subsoil and top growth yield (data pooled by selections) were improved mainly in the subsoil treatment containing the highest addition of CaCO₃.     

Characteristic responses of three tropical legumes to the inoculation of two species of VAM fungi in Andosol soils with different fertilities

The growth and mineral nutrition responses were evaluated of three tropical legumes, cowpea (Vigna unguiculata L. cv Kuromame), pigeonpea [Cajanus cajan L. (Millsp.) cv ICPL 86009] and groundnut (Arachis hypogaea cv Nakateyutaka) inoculated with two different species of VAM fungi, Glomus sp. (Glomus etunicatum-like species) and Gigaspora margarita, and grown in Andosols with different fertilities [Bray II-P: topsoil (72 ppm), subsoil (<0.1 ppm)]. Percent fungal root colonization was high in cowpea and groundnut but relatively low in pigeonpea in both soil types. Despite the low rate of root infection, significant growth responses were produced, especially in the inoculated pigeonpea plant. In all legumes, shoot dry matter production was favoured by the inoculations. Increases in shoot biomass due to mycorrhizae were greater in the subsoil than in the topsoil. Mycorrhization raised shoot concentrations of P and Ca (in cowpea and groundnut) and P and K (in pigeonpea) in the topsoil. Whereas the P concentration in shoots in the subsoil was not positively affected by VAM fungi, particularly in cowpea and pigeonpea, the concentration of K in such plants was significantly increased by VAM treatment. The results also showed that mycorrhizal enhancement of shoot micronutrient concentrations was very rare in all plants, with negative effects observed in certain cases. Cu concentration, in particular, was not affected by VAM formation in any of the plants, and Mn and Fe in pigeonpea and groundnut, respectively, remained the same whether plants were mycorrhizal or not. In both soils the three legumes responded to Glomus sp. better than to Gigaspora margarita, and the effects of the VAM fungi on each of the crops relative to the controls were greater in the subsoil than in the topsoil. However, shoot growth of groundnut was not affected as much as cowpea and pigeonpea by the type of soil used. In spite of the relatively low infection of its root, pigeonpea was generally the most responsive of the three legume species in terms of mycorrhizal/nonmycorrhizal ratios.     

The influence of rooting density in the soil on P removal by cereals on soils with different yield potential

Summary In field experiments on soils of different rooting density and different yield potential relationships between P uptake and root penetration of the soil, respectively, EUF-P contents in the topsoil were studied.On soils under uniform climatic and management conditions yields increased with increasing total root length in the subsoil (30–100 cm depth), while a negative correlation was found to exist between yields and total root length in the topsoil (0–30 cm).P removal was significantly correlated with the readily available water capacity of the rooted soil volume, which indicates the yield potential of a soil. There was no significant correlation between EUF-P contents of the soils and the P contents of the plants, but P contents in plant dry matter was significantly correlated with rooting density in the topsoil. Considering the effect on P contents of both EUF-P content in the topsoil and rooting density, the correlation coefficient was increased.     

Evaluating spatial within plot crop variability for different management practices with an optical sensor?

Subsoil constraints to root growth exacerbate frequent water and nutrient limitations to crop yields in Mediterranean-type environments. Amelioration of subsoil constraints can relieve these limitations by opening root-access to subsoil water and nutrients. However, decisions in subsoil amelioration are hampered by seasonally variable yield responses in these environments. We used the APSIM model to analyse the impact of subsoil constraints on yield and yield variability. The simulated yield data were used to calculate the financial benefits of subsoil amelioration across several scenarios. There was a strong yield-dependence on accessible soil water governed by root depth. Root depth development was limited to a minimum of either the effect of subsoil constraints or the weather-dependent depth of the soil wetting front. Insufficient rainfall in dry years or in a drier region often resulted in shallow soil wetting fronts and correspondingly shallow roots even in the absence of subsoil compaction. In these situations, there is little response to subsoil amelioration. Positive yield responses and positive financial returns to subsoil amelioration are therefore greater in good rainfall years and are more likely in a wetter region. A yield response to amelioration is also greater in coarser textured sand than finer textured sandy loam in an average rainfall season because the same amount of rainfall results in a deeper wetting front in sand than in sandy loam. Hence, roots in a sand are required to grow deeper compared to a sandy loam to access the same amount of water and therefore benefited more from subsoil amelioration in an average rainfall year. In wet years, sands leach more nitrate than sandy loam, which decreases yields and the response to subsoil amelioration in sands is more than in the sandy loam. Environmental threats occur along with yield loss when roots cannot access subsoil water. These include increased nitrate leaching and deep drainage due to unused water remaining in the soil profile. By allowing roots to access deep soil water, ameliorating subsoil is expected to yield financial gains in average to good rainfall seasons and decrease the environmental risk of drainage and leaching loss. The financial gains are expected to offset potential financial losses in dry and dry finish seasons especially in coarser textured soils and wetter environment. Responsible Editor: Jan Vos.     

Water regime of metal-contaminated soil under juvenile forest vegetation

In a three-year factorial lysimeter study in Open Top Chambers (OTCs), we investigated the effect of topsoil pollution by the heavy metals Zn, Cu, and Cd on the water regime of newly established forest ecosystems. Furthermore, we studied the influence of two types of uncontaminated subsoils (acidic vs. calcareous) and two types of irrigation water acidity (ambient rainfall chemistry vs. acidified chemistry) on the response of the vegetation. Each of the eight treatment combinations was replicated four times. The contamination (2700 mg kg^–1 Zn, 385 mg kg^–1 Cu and 10 mg kg^–1 Cd) was applied by mixing filter dust from a non-ferrous metal smelter into the upper 15 cm of the soil profile, consisting of silty loam (pH 6.5). The same vegetation was established in all 32 lysimeters. The model forest ecosystem consisted of seedlings of Norway spruce (Picea abies), willow (Salix viminalis), poplar (Populus tremula) and birch (Betula pendula) trees and a variety of herbaceous understorey plants. Systematic and significant effects showed up in the second and third growing season after canopies had closed. Evapotranspiration was reduced in metal contaminated treatments, independent of the subsoil type and acidity of the irrigation water. This effect corresponded to an even stronger reduction in root growth in the metal treatments. In the first two growing seasons, evapotranspiration was higher on the calcareous than on the acidic subsoil. In the third year the difference disappeared. Acidification of the irrigation water had no significant effect on water consumption, although a tendency to enhance evapotranspiration became increasingly manifest in the second and third year. Soil water potentials indicated that the increasing water consumption over the years was fed primarily by intensified extraction of water from the topsoil in the lysimeters with acidic subsoil, whereas also lower depths became strongly exploited in the lysimeters with calcareous subsoil. These patterns agreed well with the vertical profiles of fine root density related with the two types of subsoil. Leaf transpiration measurements and biomass samples showed that different plant species in part responded quite differently and occasionally even in opposite ways to the metal treatments and subsoil conditions. They suggest that the year-to-year changes in treatment effects on water consumption and extraction patterns were related to differences in growth dynamics, as well as to shifts in competitiveness of the various species. Results showed that the uncontaminated subsoil offered a possibility to compensate the reduction in root water extraction in the topsoil under drought, as well as metal stress.     

RFLP mapping of manganese efficiency in barley

In many cropping regions of the world, yield is limited by the availability of micronutrients, and micronutrient-efficient cultivars provide a yield advantage. Traditional methods of testing cultivars for micronutrient efficiency are time-consuming and laborious. Molecular markers linked to loci controlling micronutrient efficiency will allow more rapid and efficient selection and introgression of these traits than is currently possible. Using a pot-based bioassay and bulked segregant analysis of an F₂ population, we have identified several RFLPs (grouped distally on chromosome 4HS) linked to a locus for manganese efficiency in barley. This manganese efficiency locus has been designated Mel1. Pot bioassay analysis of intercrosses suggests that three useful sources of manganese efficiency are likely to be allelic at the Mel1 locus. Field evaluation of marker selected F₄ progeny supports the major role of Mel1 in the genetic control of manganese efficiency. Adoption of marker assisted selection for this trait in the Southern Australian barley breeding program has occurred. This has been facilitated by the demonstration that the Mel1 allele of Amagi Nijo can be distinguished from 95 other locally useful varieties and breeder’s lines on the basis of RFLPs identified by just two molecular markers. Received: 20 October 1999 / Accepted: 18 February 2000     

藏东南典型暗针叶林不同土壤剖面微生物群落特征

深层土壤中的微生物群落对陆地生态系统养分和能量循环转化过程不可或缺,研究青藏高原典型暗针叶林带土壤微生物群落在土壤垂直剖面的变化特征,对深入认识高寒区域森林生态系统土壤微生物群落构建特征及全球变化影响预测具有重要意义。运用Illumina Miseq高通量测序技术和分子生态网络分析,研究藏东南色季拉山暗针叶林带表层（0-20 cm）和底层土壤（40-60 cm）微生物群落组成及分子生态网络结构。研究结果表明随着土壤深度增加,真菌和细菌的丰富度和Shannon多样性指数显著降低。主坐标分析（PCoA）显示土壤深度显著影响真菌和细菌的群落结构（P < 0.01）。不同微生物种群对土壤深度的响应有显著差异,座囊菌纲（Dothideomycetes）、银耳纲（Tremellomycetes）和拟杆菌门（Bacteroidetes）、变形菌门（Proteobacteria）的相对丰度随剖面加深而显著降低,而古菌根菌纲（Archaeorhizomycetes）和绿弯菌门（Chloroflexi）则显著增加。分子生态网络分析发现,真菌网络以负相关连接为主（占总连接数65%-98%）,而细菌网络以正相关连接为主（69%-75%）,真菌和细菌网络中正相关连接的比例均随剖面加深而增加。底层土壤真菌和细菌网络的平均连接度和平均聚类系数均高于表层土壤,说明微生物网络随土壤深度的增加而变得更复杂。真菌网络的平均路径距离和模块性在底层土壤均大于表层土壤,意味着真菌网络应对环境变化的稳定性随剖面加深而增加,而细菌网络则正相反,在表层土壤的稳定性更强。真菌网络中连接节点的个数随剖面加深而增加,锤舌菌纲（Leotiomycetes）是连接网络模块的关键菌种;在细菌网络中模块枢纽和连接节点则随剖面加深而降低,并且放线菌门、变形菌门等关键种群在分子生态网络中的功能在表层和底层土壤有明显差异。综上所述,藏东南色季拉山暗针叶林带深层土壤中微生物群落特征与表层土壤有显著差别,揭示影响深层土壤微生物网络构建和稳定的关键种群,对深入理解和预测青藏高原森林生态系统对全球变化的响应与反馈有重要意义。     

Root growth of triticale and barley grown for grain or for forage-plus-grain in a Mediterranean climate

Root systems of one triticale (× triticosecale Witt.) and one barley (Hordeum vulgare L.) cultivar grown for grain or for the dual purpose of winter forage-plus-grain were studied in a Mediterranean climate (Granada, Spain). The aim was to assess the effect of winter forage removal on root systems and to improve the knowledge of cereal root systems under Mediterranean conditions in relation to soil water use. After the forage was removed by clipping at the end of the winter period, cereal roots were shallower and lower in length densities, compared to the unclipped treatment. The largest differences occurred during the clipping-anthesis period and in the upper soil layers. At the end of the life cycle, the differences between the two systems regarding depth, density and dry matter of roots were small or nil. Moreover, there were no differences in total water use between clipped and unclipped cereals.Under both production systems (grain and forage plus grain), cereals demonstrated variable downward root extension (0.9 to 1.8 m) as a response to the wetting depth. Triticale roots continued growing after anthesis, especially in the deeper soil layers. In spite of that, root systems were not able to extract a notable amount of residual water (25 to 50 mm) apparently available from the subsoil. In semi-arid Mediterranean drylands, cereal root systems with greater phenotypic plasticity (deeper or larger in the subsoil) in response to subsoil water should be of interest in wetter areas or seasons. This does not necessarily imply a larger root system, but rather a root growth pattern with greater root growth in the subsoil.     

The effect of soil water status on fertiliser, topsoil and subsoil phosphorus utilisation by wheat

Background and aims

Crop phosphorus (P) content is controlled by P uptake from both banded P fertiliser and from P throughout the soil profile. These P supply factors are in turn controlled by soil, climatic and plant factors. The aim of this experiment was to measure the contribution of fertiliser, topsoil and subsoil P to wheat plants under wet and dry growing season conditions.

Methods

An isotopic tracer technique was used to measure P uptake from fertiliser at seven agricultural field sites under wet and dry growing season conditions. At three of these sites a dual isotopic technique was used to distinguish between wheat uptake of P from fertiliser, topsoil (0–15 cm) and subsoil (below 15 cm).

Results

The amount of P fertiliser used by wheat was in the order of 3–30% of the P applied and increased with increasing rainfall. Topsoil P was the most important P source, but when sufficient P was present in the subsoil, P fertiliser addition stimulated the use of subsoil P.

Conclusions

Most crop P uptake was from the topsoil, however P fertiliser banded below the seed increased plant P uptake and stimulated the use of subsoil P in one soil type in a decile 7 (above average rainfall) growing season.     

Tolerance and avoidance of Al toxicity by Mucuna pruriens var. utilis at different levels of P supply

Previous laboratory experiments showed that velvet bean Mucuna pruriens is moderately tolerant to the presence of Al (up to 185 µM) in the root environment, but that it only develops a shallow root system in acid soils. Field experiments showed that Mucuna can tolerate acid subsoil conditions in a homogeneous root environment, but avoids subsoil if topsoil is present. Subsequent split-root experiments with a recirculating nutrient solution showed that this subsoil avoidance may be based on an Al avoidance mechanism in the root system. This Al avoidance mechanism, however, was not evident when phosphorus (P) supply to the whole plant was adequate. We thus hypothesized that surface application of P may help to overcome Al avoidance in the subsoil.In a field experiment on an ultisol in Lampung (Indonesia), only a moderate increase in aboveground biomass production was found for a wide range of P application rates, although the soil was low in available P, and the P adsorption isotherm was very steep. An increased P status of the topsoil and an increased P concentration in the aboveground biomass (from 50 to 75 mmol kg^-1) had no effect on root development in the subsoil.     

Subsoil root activity in tree-based cropping systems

An increasing number of studies indicate that (i) nutrient and water resources can be abundant in the subsoil and (ii) trees have deep root systems that can possibly reach these resources. It is less clear whether subsoil resources are actually improving water and nutrient status of tree-based cropping systems and whether they are significantly increasing crop production and yield. To answer such a question, the distribution of nutrient and water uptake by trees needs to be quantified. So-called `root activity distributions' give valuable information about actual subsoil use by trees whereas studies on root length or mass distributions do not often correlate with uptake distributions. Despite the usually lower relative root activity in the subsoil compared to the topsoil per unit soil, the large volume of subsoil in comparison to mostly shallow topsoil is an important resource for crop nutrient and water uptake. The present study compares published root activity distributions using the model Activity=A _max(1–k ^depth). The obtained regression constants k of 0.91–0.99 determined in this publication reflect the values computed by an earlier published survey for root biomass ranging from tundra to those of temperate forest biomes. Thus, tree crops can have shallow root activity and 75% of their total root activity in the first 0.1 m of soil, or very deep root activity with more than 90% below 0.1 m. Neither environmental factors (i.e., climate and soil properties available from these publications) nor plant species explain differences of root activity distributions with depth. The deepest root activity is found for fruit trees such as citrus, guava and mango. Shaded crops such as coffee and cacao tend to have shallower root activity than fruit trees. Monocots including oil palm, coconut or banana have root activity that can be both deep and shallow. Regional and temporal variations of subsoil root activity for the same tree species are significant and generally larger than differences between species. Root activity patterns of tree crops appear to be sufficiently flexible to allow for subsoil resource use. Consequently, management such as pruning, fertilization, liming and irrigation are shown to significantly affect subsoil root activity.     

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.