Management of biological N2 fixation in alley cropping systems: Estimation and contribution to N balance

Alley cropping is being widely tested in the tropics for its potential to sustain adequate food production with low agricultural inputs, while conserving the resource base. Fast growth and N yield of most trees used as hedgerows in alley cropping is due greatly to their ability to fix N₂ symbiotically with Rhizobium. Measurements of biological N₂ fixation (BNF) in alley cropping systems show that some tree species such as Leucaena leucocephala, Gliricidia sepium and Acacia mangium can derive between 100 and 300 kg N ha^-1 yr^–1 from atmospheric N₂, while species such as Faidherbia albida and Acacia senegal might fix less than 20 kg N ha^-1 yr^-1. Other tree species such as Senna siamea and S. spectabilis are also used in alley cropping, although they do not nodulate and therefore do not fix N₂. The long-term evaluation of the potential or actual amounts of N₂ fixed in trees however, poses problems that are associated with their perennial nature and massive size, the great difficulty in obtaining representative samples and applying reliable methodologies for measuring N₂ fixed. Strategies for obtaining representative samples (as against the whole tree or destructive plant sampling), the application of ¹⁵N procedures and the selection criteria for appropriate reference plants have been discussed.Little is known about the effect of environmental factors and management practices such as tree cutting or pruning and residue management on BNF and eventually their N contribution in alley cropping. Data using the ¹⁵N labelling techniques have indicated that up to 50% or more of the tree's N may be below ground after pruning. In this case, quantification of N₂ fixed that disregards roots, nodules and crowns would result in serious errors and the amount of N₂ fixed may be largely underestimated. Large quantities of N are harvested with hedgerow prunings (>300 kg N ha^-1 yr^-1) but N contribution to crops is commonly in the range of 40–70 kg N ha^-1 season. This represents about 30% of N applied as prunings; however, N recoveries as low as 5–10% have been reported. The low N recovery in maize (Zea mays) is partly caused by lack of synchronization between the hedgerow trees N release and the associated food crop N demand. The N not taken up by the associated crop can be immobilized in soil organic matter or assimilated by the hedgerow trees and thus remain in the system. This N can also be lost from the system through denitrification, volatilization or is leached beyond the rooting zone. Below ground contribution (from root turnover and nodule decay) to an associated food crop in alley cropping is estimated at about 25–102 kg N ha^-1 season^-1. Timing and severity of pruning may allow for some management of underground transfer of fixed N₂ to associated crops. However many aspects of root dynamics in alley cropping systems are poorly understood. Current research projects based on ¹⁵N labelling techniques or ¹⁵N natural abundance measurements are outlined. These would lead to estimates of N₂ fixation and N saving resulting from the management of N₂ fixation in alley cropping systems.     

Decomposition and stabilization of root litter in top- and subsoil horizons: what is the difference?

Mechanisms leading to high mean residence times of organic matter in subsoil horizons are poorly understood. In lower parts of the soil profile root material contributes greatly to soil organic matter (SOM). The objective of this study was to elucidate the decomposition dynamics of root-derived C and N in different soil depths during a 3 year field experiment and to examine the importance of different protection mechanisms as well as abiotic factors for the decomposition dynamics. Additionally, we assessed the effect of root litter addition on native SOM. Our conceptual approach included the exposure of litterbags with ¹³C and ¹⁵N labeled wheat root material mixed to loamy agricultural soil at three different soil depths (30, 60 and 90 cm). During the incubation period, we monitored soil temperature, humidity and the incorporation of root derived C and N into the soil microbial biomass and physical SOM fractions. Our results showed that abiotic decay conditions were better in subsurface horizons compared to the topsoil. Root litter addition significantly increased the size of microbial biomass in all three soil horizons. In the topsoil, root-derived C decomposition was significantly higher in the first 6 months of incubation compared to subsoil horizons. In 60 and 90 cm depths, a lag phase with development of soil microbial biomass seemed to be prevailing before decomposition was activated. For root-derived N, similar decomposition kinetics could be observed in top- and subsoil horizons. Despite of higher SOM contents, better soil structure and higher microbial activity in the topsoil horizon compared to subsoil horizons, the amounts of root-derived C and N remaining after 3 years were similar for all three depths. Most of the root-derived C and N was present as organo-mineral complexes or occluded in soil aggregates (oPOM), illustrating similar importance of these two protection mechanisms in all three soil depths. Addition of fresh root litter caused small losses of native soil C whereas native N was retained. We conclude that despite of similar SOM protection mechanisms, there are distinct differences in decomposition dynamics of root litter between top- and subsoil horizons. In the long run, the better abiotic decay conditions prevailing in subsoil horizons may compensate for their poorer physico-chemical characteristics.     

Influence of pruning management on P and N distribution and use efficiency by N2 fixing and non-N2 fixing trees used in alley cropping systems

Pruning of hedgerow trees is an important management practice for the successful establishment of an alley cropping system. Although pruning affects biomass production, only meager evidence of this management on distribution of nutrients among the different plant organs after tree regrowth is available. This study examined the effect of pruning on the distribution and use efficiency of N and P in a N₂ fixing leguminous tree species, Gliricidia sepium, and two non-N₂ fixing leguminous tree species, Senna siamea and S. spectabilis, grown in a field on an Alfisol (low in P) at Fashola (Guinea Savanna Zone), Southwestern Nigeria. Four P rates, 0, 20, 40 and 80 kg P ha^–1 as single superphosphate were used and management treatments included pruned versus unpruned plants. The ¹⁵N isotope dilution technique was used to measure N₂ fixation in G. sepium. Partitioning of total P among different plant organs was influenced by plant species and pruning management, but was not affected by P application rates. The distribution of total P in the various plant organs followed that of dry matter yield while N partitioning had a different pattern. Pruned plants distributed about 118% more total P to branches and had a higher physiological P use efficiency (PPUE) than unpruned plants. Leaves were the biggest sink for total N and N allocation in the other plant organs was influenced by plant species and pruning management, G. sepium had relatively more of its total N and P partitioned into roots (about double that of the non-N₂ fixing trees) but had a lower PPUE. Unpruned and pruned G. sepium derived 35 and 54% respectively of their total N from atmospheric N2, with about 54% of the fixed N2 being allocated to leaves and roots. Results showed that N and P pools turned over in the branches during plant regrowth after pruning but the causative factors associated with this phenomenon were not clear.     

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.     

Decomposition of leaves of huisache (Acacia tortuoso) and mesquite (Prosopis spp) in soil of the central highlands of Mexico

In the central highlands of Mexico, mesquite (Prosopis spp) and huisache (Acacia tortuoso), N₂ fixing trees or shrubs, dominate the vegetation and are used in an alley cropping system to prevent erosion and restore soil fertility. We investigated how much the leaves of both trees contribute to dynamics of carbon (C) and nitrogen (N) in soil by adding leaves of both species to soil sampled under the canopy of mesquite and huisache, outside their canopy and from fields cultivated with maize at three different sites and monitoring microbial biomass C, production of carbon dioxide (CO₂), and dynamics of inorganic N (ammonium and nitrate) in an aerobic incubation. The soluble fraction and N content of the mesquite leaves were larger than in the huisache leaves, but lignin and polyphenol content were lower. Evolution of CO₂ increased 2.7-times when mesquite and 2.4-times when huisache leaves were added to soil. During all stages of decomposition and in all treatments, C mineralization of leaves from mesquite was greater than from huisache leaves. Mesquite leaves induced an increase in mineral N of 25.6 mg N kg^–1 soil after 56 days and those of huisache 9.8 mg N kg^–1. Twenty-six percent of N from mesquite leaves and 11% of huisache was mineralized, if no priming effect was considered. Nitrogen release from the leaves was greater when the soil organic matter content was lower. It was found that soil under the canopy of mesquite and huisache effectively accumulated organic material, micro-organisms and valuable nutrients. In an alley cropping system huisache might be a better choice than mesquite as huisache grows faster than mesquite and sheds its leaves twice a year while mesquite only once, although the amount of N mineralized was larger from mesquite leaves than from those of huisache.     

Carbon and nitrogen in a temperate agroforestry system: Using stable isotopes as a tool to understand soil dynamics

Natural exchanges of carbon (C) between the atmosphere, the oceans, and terrestrial ecosystems are currently being modified through human activities as a result of fossil fuel burning and the conversion of tropical forests to agricultural land. These activities have led to a steady increase of atmospheric carbon dioxide (CO₂) over the last two Centuries. The goal of this study was to determine the potential of temperate agroforestry systems to sequester C in soil. Therefore, changes in the soil organic C (SOC) and nitrogen (N) pools were quantified and the δ¹³C and δ¹⁵N stable isotope technique was applied to assess soil C and N dynamics in a 13-year old hybrid poplar alley cropping system in Southern Canada. Results from this study showed that after 13 years of alley cropping the SOC and N pools did not differ significantly (p = 0.01) with distance from the tree row, although a trend of a larger SOC and N pool near the tree row could be observed. Soil organic C after 13 years of alley cropping, was 19 mg C g⁻¹ compared to 11 mg C g⁻¹ upon initiation of agroforestry. Soil organic C and N were not evenly distributed throughout the plow layer. The largest C and N pool occurred in the top 20 cm, which is due to the accumulation of organic material in the upper horizons as a result of no-till cultivation. The entire soil, to a 40 cm depth, showed a δ¹³C shift to that of C₃ residue. This shift reflects the greater input of residues from C₃ plants such as that derived from beans, wheat, and hybrid poplar leaf litterfall. The proportion of C derived from a C₃ source ranged from 64 to 69% to a 40 cm depth. The soil δ¹⁵N signature of this study is similar to that of mineral soil, and reflect values characteristic of N mineralization processes. However, the entire soil shows a positive shift in δ¹⁵N as a result of historical additions of manure and current use of mineral fertilizers, and ongoing processes of denitrification and nitrate leaching, which leads to an enrichment of the soil.     

Incorporation of shoot versus root-derived <Superscript>13</Superscript>C and <Superscript>15</Superscript>N into mineral-associated organic matter fractions: results of a soil slurry incubation with dual-labelled plant material

Mineral-associated organic matter (MAOM) is a key component of the global carbon (C) and nitrogen (N) cycles, but the processes controlling its formation from plant litter are not well understood. Recent evidence suggests that more MAOM will form from higher quality litters (e.g., those with lower C/N ratios and lower lignocellulose indices), than lower quality litters. Shoots and roots of the same non-woody plant can provide good examples of high and low quality litters, respectively, yet previous work tends to show a majority of soil organic matter is root-derived. We investigated the effect of litter quality on MAOM formation from shoots versus roots using a litter-soil slurry incubation of isotopically labeled (¹³C and ¹⁵N) shoots or roots of Big Bluestem (Andropogon gerardii) with isolated silt or clay soil fractions. The slurry method minimized the influence of soil structure and maximized contact between plant material and soil. We tracked the contribution of shoot- and root-derived C and N to newly formed MAOM over 60 days. We found that shoots contributed more C and N to MAOM than roots. The formation of shoot-derived MAOM was also more efficient, meaning that less CO₂ was respired per unit MAOM formed. We suggest that these results are driven by initial differences in litter chemistry between the shoot and root material, while results of studies showing a majority of soil organic matter is root-derived may be driven by alternate mechanisms, such as proximity of roots to mineral surfaces, greater contribution of roots to aggregate formation, and root exudation. Across all treatments, newly formed MAOM had a low C/N ratio compared to the parent plant material, which supports the idea that microbial processing of litter is a key pathway of MAOM formation.     

Tree species selection and soil tillage in alley cropping systems with Phaseolus vulgaris L. in a humid premontane climate: biomass production,nutrient cycling and crop responses

The development of alley cropping systems is based on the assumption that leguminous trees planted in hedgerows influence the yield of associated crops favourably by means of the additional nutrient pool applied to the soil through tree prunings. An on-station field study (split-plot design in a randomised block design) was conducted on an Eutric Cambisol under humid premontane climate conditions in Costa Rica in order to evaluate the ability of Erythrina poeppigiana, Calliandra calothyrsus and Gliricidia sepium to increase bean (Phaseolus vulgaris) yields compared to sole cropping. Soil tillage was applied as a sub-treatment in order to evaluate if soil preparation would additionally alter soil fertility and bean yield. After seven years with pruning twice per year, the size of both the total N and P pool in the pruned tree material was about three times higher for Erythrina prunings than for Calliandra and Gliricidia prunings. Two and five weeks after mulch application 50–150% higher inorganic N pools were measured in the soil from Erythrina plots, the bean shoot biomass at harvest was increased by 65–100% and the bean yield was 15–50% higher than in plots with beans alone. Hence, of the three tree species, Erythrina was the best choice for alley cropping systems in the pedoclimatic environment studied. Soil tillage reduced bean yield, soil organic matter, total soil N content and soil microbial biomass N in the top soil and is not recommended for similar soils in humid premontane climates.     

Alley cropping with Senna siamea in South-western Nigeria: II. Dry matter,total N,and urea-derived N dynamics of the Senna and maize roots

Crop and tree roots are crucial in the nutrient recycling hypotheses related to alley cropping systems. At the same time, they are the least understood components of these systems. The biomass, total N content and urea-derived N content of the Senna and maize roots in a Senna-maize alley cropping system were followed for a period of 1.5 years (1 maize-cowpea rotation followed by 1 maize season) to a depth of 90 cm, after the application of ¹⁵N labeled urea. The highest maize root biomass was found in the 0–10 cm layer and this biomass peaked at 38 and 67 days after planting the 1994 maize (DAP) between the maize rows (112 kg ha^–1, on average) and at 38, 67 and 107 DAP under the maize plants (4101 kg ha^–1, on average). Almost no maize roots were found below 60 cm at any sampling date. Senna root biomass decreased with time in all soil layers (from 512 to 68 kg ha^–1 for the 0–10 cm layer between 0 and 480 DAP). Below 10 cm, at least 62% of the total root biomass consisted of Senna roots and this value increased to 87% between 60 and 90 cm. Although these observations support the existence of a Senna root `safety net' between the alleys which could reduce nutrient leaching losses, the depth of such a net may be limited as the root biomass of the Senna trees in the 60–90 cm layer was below 100 kg ha^–1, equivalent to a root length density of only < 0.05 cm cm^–3. The proportion of maize root N derived from the applied urea (%Ndfu) decreased significantly with time (from 21% at 21 DAP to 8% at 107 DAP), while %Ndfu of the maize roots at the second harvest (480 DAP) was only 0.6%. The %Ndfu of the Senna roots never exceeded 4% at any depth or sampling time, but decreased less rapidly compared to the %Ndfu of the maize roots. The higher %Ndfu of the maize roots indicates that maize is more efficient in retrieving urea-derived N. The differences in dynamics of the %Ndfu also indicate that the turnover of N through the maize roots is much faster than the turnover of N through the Senna roots. The recovery of applied urea-N by the maize roots was highest in the top 0–10 cm of soil and never exceeded 0.4% (at 38 DAP) between the rows and 7.1% (at 67 DAP) under the rows. Total urea N recovery by the maize roots increased from 1.8 to 3.2% during the 1994 maize season, while the Senna roots never recovered more than 0.8% of the applied urea-N at any time during the experimental period. These values are low and signify that the roots of both plants will only marginally affect the total recovery of the applied urea-N. Measurement of the dynamics of the biomass and N content of the maize and Senna roots helps to explain the observed recovery of applied urea-N in the aboveground compartments of the alley cropping system.     

The nitrogen handling characteristics of white clover (Trifolium repens L.) cultivars and a perennial ryegrass (Lolium perenne L.) cultivar

Ryegrass (Lolium perenne L.) and white clover (Trifolium repens L.) have contrasting responses to soil mineral N availability and clover has the ability to fix atmospheric N(2) symbiotically. It has been hypothesized that these differences are the key to understanding grass-clover coexistence and vegetative dynamics in pastures. However, the whole plant response of clover and ryegrass to mineral N availability has not been fully characterized and inter-cultivar variability in the N-handling dynamics of clover has not been assessed. A detailed experimental study to address these issues was undertaken. For all clover cultivars and ryegrass, mass specific mineral N uptake rates (of whole plants) were similar saturating functions of mineral N availability. For all clover cultivars total N assimilation rates, whole plant C : N ratios and root : shoot ratios were independent of mineral N availability. Clover growth rates were also independent of mineral N availability except for a slight (<10%) reduction at very low N availability levels. Specific N(2) fixation rate (whole plant) was precisely controlled to ensure fixation balanced the deficit between mineral N uptake and the total N assimilation required to maintain constant whole plant C : N ratio. There was always a deficit between N uptake and the total N assimilation required to maintain C : N ratio. Consequently, some N(2) fixation remained engaged even at high mineral N availability levels. All inter-cultivar variation in N(2) fixation dynamics could be attributed to variations in growth rate. Clover mass specific growth rate declined as plant size increased. Ryegrass specific growth rate, whole plant C : N ratio and root : shoot ratio were dependent on N availability. Increased N availability led to increased growth rate and decreased C : N and root : shoot ratios. Specific growth rate was also dependent on plant size, growth rate declining as plant size increased. It is concluded that clover inter-cultivar variation in field performance is unlikely to be a consequence of variation in N-handling characteristics. Inter-cultivar differences in growth rate are likely to be a much more important source of variation.     

模拟根系分泌物C:N化学计量特征对川西亚高山森林土壤碳动态和微生物群落结构的影响

目前有关森林根系分泌物及其诱导的土壤生态学效应研究主要关注根系碳(C)源输入, 而极少关注根系分泌物氮(N)源输入及其伴随的C:N化学计量特征对土壤过程和功能的影响, 极大地限制了我们对森林根系-土壤-微生物互作机制的深入认识。该研究以川西亚高山天然林和云杉(Picea asperata)人工林土壤为对象, 模拟配制不同C:N化学计量特征(只有N、C:N = 10、C:N = 50、C:N = 100和只有C处理)的根系分泌物溶液进行人工添加试验, 以探究根系分泌物化学计量特征对两种林分土壤碳动态及其微生物群落结构的影响差异。结果表明: 模拟根系分泌物C添加总体促进了两种林分土壤有机质分解激发效应而降低了土壤总碳(TC)含量, 而N添加在一定程度上缓和了两种林分土壤TC含量的降低幅度, 且C添加导致天然林土壤TC含量的降低幅度明显低于土壤N有效性更低的人工林。几种根系分泌物添加处理对两种林分土壤活性和惰性碳库的影响无明显规律。另外, 根系分泌物C添加总体降低了天然林土壤微生物总磷脂脂肪酸(PLFA)含量和细菌、放线菌、真菌PLFA含量, 而总体增加人工林土壤微生物PLFA总量和细菌、放线菌、真菌PLFA含量, 并诱导两种林分土壤微生物群落结构(细菌:真菌相对丰度)也发生了各自不同的变化。上述结果表明森林根系分泌物N源输入和土壤N有效性共同调控根系C源输入对土壤有机质分解激发效应的方向和幅度。研究结果为深入揭示典型森林根系分泌物化学计量特征对土壤生物化学循环过程的调控机制提供了一定的理论依据。     

Combined effects of atmospheric CO<Subscript>2</Subscript> and N availability on the belowground carbon and nitrogen dynamics of aspen mesocosms

It is uncertain whether elevated atmospheric CO₂ will increase C storage in terrestrial ecosystems without concomitant increases in plant access to N. Elevated CO₂ may alter microbial activities that regulate soil N availability by changing the amount or composition of organic substrates produced by roots. Our objective was to determine the potential for elevated CO₂ to change N availability in an experimental plant-soil system by affecting the acquisition of root-derived C by soil microbes. We grew Populus tremuloides (trembling aspen) cuttings for 2 years under two levels of atmospheric CO₂ (36.7 and 71.5 Pa) and at two levels of soil N (210 and 970 μg N g^–1). Ambient and twice-ambient CO₂ concentrations were applied using open-top chambers, and soil N availability was manipulated by mixing soils differing in organic N content. From June to October of the second growing season, we measured midday rates of soil respiration. In August, we pulse-labeled plants with ¹⁴CO₂ and measured soil ¹⁴CO₂ respiration and the ¹⁴C contents of plants, soils, and microorganisms after a 6-day chase period. In conjunction with the August radio-labeling and again in October, we used ¹⁵N pool dilution techniques to measure in situ rates of gross N mineralization, N immobilization by microbes, and plant N uptake. At both levels of soil N availability, elevated CO₂ significantly increased whole-plant and root biomass, and marginally increased whole-plant N capital. Significant increases in soil respiration were closely linked to increases in root biomass under elevated CO₂. CO₂ enrichment had no significant effect on the allometric distribution of biomass or ¹⁴C among plant components, total ¹⁴C allocation belowground, or cumulative (6-day) ¹⁴CO₂ soil respiration. Elevated CO₂ significantly increased microbial ¹⁴C contents, indicating greater availability of microbial substrates derived from roots. The near doubling of microbial ¹⁴C contents at elevated CO₂ was a relatively small quantitative change in the belowground C cycle of our experimental system, but represents an ecologically significant effect on the dynamics of microbial growth. Rates of plant N uptake during both 6-day periods in August and October were significantly greater at elevated CO₂, and were closely related to fine-root biomass. Gross N mineralization was not affected by elevated CO₂. Despite significantly greater rates of N immobilization under elevated CO₂, standing pools of microbial N were not affected by elevated CO₂, suggesting that N was cycling through microbes more rapidly. Our results contained elements of both positive and negative feedback hypotheses, and may be most relevant to young, aggrading ecosystems, where soil resources are not yet fully exploited by plant roots. If the turnover of microbial N increases, higher rates of N immobilization may not decrease N availability to plants under elevated CO₂. Received: 12 February 1999 / Accepted: 2 March 2000     

Estimating belowground nitrogen inputs of pea and canola and their contribution to soil inorganic N pools using 15N labeling

Background and aims

Crop species grown in a diversified crop rotation can influence soil N dynamics to varying degrees due to differences in the quantity and quality of the residues returned to the soil. The aim of this study was to quantify the contribution of N rhizodeposition by canola (Brassica napus L.) and pea (Pisum sativum L.) to the crop residue N balance and soil inorganic N pool.

Methods

Canola and pea were grown in a soil-sand mixture and were subject to cotton-wick ¹⁵N labeling in a greenhouse experiment. Nitrogen-15 recovered in the soil and roots were used to estimate N rhizodeposition.

Results

Belowground N, including root N and N rhizodeposits, comprised 70 % and 61 % of total crop residue N for canola and pea, respectively. Canola released the greatest amount of total root-derived N to the soil, which was related to greater root biomass production by canola. However, root-derived N in the soil inorganic N pool was greater under pea (13 %) than canola (4 %).

Conclusions

Our results show a significant belowground N contribution to total crop residue from pea and canola. Further investigation is required to determine whether input of the more labile N rhizodeposits of pea improves soil N supply to succeeding crops or increases the potential for N loss from the soil system relative to canola.     

Indirect Effects of Nitrogen Amendments on Organic Substrate Quality Increase Enzymatic Activity Driving Decomposition in a Mesic Grassland

The fate of soil organic carbon (SOC) is determined, in part, by complex interactions between the quality of plant litter inputs, nutrient availability, and the microbial communities that control decomposition rates. This study explores these interactions in a mesic grassland where C and nitrogen (N) availability and plant litter quality have been manipulated using both fertilization and haying for 7 years. We measured a suite of soil parameters including inorganic N, extractable organic C and N (EOC and EON), soil moisture, extracellular enzyme activity (EEA), and the isotopic composition of C and N in the microbial biomass and substrate sources. We use these data to determine how the activity of microbial decomposers was influenced by varying levels of substrate C and N quality and quantity and to explore potential mechanisms explaining the fate of enhanced plant biomass inputs with fertilization. Oxidative EEA targeting relatively recalcitrant C pools was not affected by fertilization. EEA linked to the breakdown of relatively labile C rich substrates exhibited no relationship with inorganic N availability but was significantly greater with fertilization and associated increases in substrate quality. These increases in EEA were not related to an increase in microbial biomass C. The ratio of hydrolytic C:N acquisition enzymes and δ¹³C and δ¹⁵N values of microbial biomass relative to bulk soil C and N, or EOC and EON suggest that microbial communities in fertilized plots were relatively C limited, a feature likely driving enhanced microbial efforts to acquire C from labile sources. These data suggest that in mesic grasslands, enhancements in biomass inputs and quality with fertilization can prompt an increase in EEA within the mineral soil profile with no significant increases in microbial biomass. Our work helps elucidate the microbially mediated fate of enhanced biomass inputs that are greater in magnitude than the associated increases in mineral soil organic matter.     

Nitrogen cycling in leucaena (Leucaena leucocephala) alley cropping in semi-arid tropics

In an alley cropping system, prunings from the hedgerow legume are expected to supply nitrogen (N) to the associated cereal. However, this may not be sufficient to achieve maximum crop yield. Three field experiments with alley-cropped maize were conducted in a semi-arid environment in northern Australia to determine: (1) the effect of N fertilizer on maize growth in the presence of fresh leucaena prunings; (2) the effect of incorporation of leucaena and maize residues on maize yield and the fate of plant residue¹⁵N in the alley cropping system; and (3) the¹⁵N recovery by maize from¹⁵N-labelled leucaena, maize residues and ammonium sulphate fertilizer.Leucaena residues increased maize crop yield and N uptake although they did not entirely satisfy the N requirement of the alley crop. Additional N fertilizer further increased the maize yield and N uptake in the presence of leucaena residues. Placement of leucaena residues had little effect on the availability of N to maize plants over a 2 month period. The incorporation of leucaena residues in the soil did not increase the recovery of leucaena¹⁵N by maize compared with placement of the residues on the soil surface. After 2 months, similar proportions of the residue¹⁵N were recovered by maize from mulched leucaena (6.3%), incorporated leucaena (6.1%) and incorporated maize (7.6%). By the end of one cropping season (3 months after application) about 9% of the added¹⁵N was taken up by maize from either¹⁵N-labelled leucaena as mulch or¹⁵N-labelled maize residues applied together with unlabelled fresh leucaena prunings as mulch. The recovery of the added¹⁵N was much higher (42.7%) from the¹⁵N-labelled ammonium sulphate fertilizer at 40 kg N ha^-1 in the presence of unlabelled leucaena prunings. Most of the added¹⁵N recovered in the 200 cm soil profile was distributed in the top 25 cm soil with little leached below that. About 27–41% of the leucaena¹⁵N was apparently lost, largely through denitrification from the soil and plant system, in one cropping season. This compared with 35% of the fertilizer¹⁵N lost when the N fertilizer was applied in the presence of prunings. ei]H Lambers     

Short-term incorporation of freshly fixed plant carbon into the soil animal food web: field study in a spruce forest

We analyzed the dynamics of the short-term incorporation of recently fixed carbon into the below-ground food web in a boreal forest. Five young spruce trees (Picea abies) were pulse-labeled with ¹³CO₂ and the isotopic label was traced in soil invertebrates during 5 weeks. The freshly fixed plant carbon quickly entered both litter-located and soil-located compartments of the detrital food web. Among invertebrates inhabiting the mineral soil layers, a trophic link to the root-derived C was most pronounced in species with higher δ ¹⁵N values, suggesting this energy source to be more important in deeper mineral soil horizons. The label appeared faster in saprophagous animals than in predators (the median time lag after labeling was 6 and 12 days, respectively), but the difference was not significant. The label was recovered in 15 of 38 species of saprophagous animals and in 20 of 63 species of predators. Among saprophages, the frequency and intensity of the label was relatively high in endogeic collembolans and in bibionid larvae, but earthworms and enchytraeids were not labeled. Several groups of predators, lithobiid centipedes in particular, quickly acquired the root-derived carbon, possibly indicating the feeding on live roots or mycorrhizal mycelium. In total, only 35 % of species or genera examined acquired the label. This suggests that majority of invertebrate taxa in the decomposer food web are unlikely to depend heavily on freshly fixed plant carbon provided by roots and root-associated microorganisms.     

Spatial distribution and turnover of root-derived carbon in alfalfa rhizosphere depending on top- and subsoil properties and mycorrhization

Aims

This study analyzed the extent to which root exudates diffuse from the root surface towards the soil depending on topsoil and subsoil properties and the effect of arbuscular mycorrhizal fungal hyphae on root-derived C distribution in the rhizosphere.

Methods

Alfalfa was grown in three-compartment pots. Nylon gauze prevented either roots alone or roots and arbuscular mycorrhizal fungal hyphae from penetrating into the rhizosphere compartments. ¹⁴CO₂ pulse labeling enabled the measurement of ¹⁴C-labeled exudates in dissolved (DOC) and total organic carbon (TOC) in the rhizosphere, distributed either by diffusion alone or by diffusion, root hair and hyphal transport.

Results

Root exudation and microbial decomposition of exudates was higher in the rhizosphere with topsoil compared to subsoil properties. Exudates extended over 28 mm (DOC) and 20 mm (TOC). Different soil properties and mycorrhization, likely caused by the low arbuscular mycorrhizal colonization of roots (13?±?4 % (topsoil properties) and 18?±?5 % (subsoil properties)), had no effect.

Conclusions

Higher microbial decomposition compensated for higher root exudation into the rhizosphere with topsoil properties, which resulted in equal exudate extent when compared to the rhizosphere with subsoil properties. Higher ¹⁴C activity used for labeling compared with previous studies enabled the detection of low exudate concentrations at longer distances from the root surface.     

Phosphorus availability under alley cropping and mulched and unmulched sole cropping systems in Costa Rica

Phosphorus availability was measured in soils under five cropping systems: alley cropping with Erythrina poeppigiana, alley cropping with Gliricidia sepium, sole cropping with Erythrina poeppigiana mulch applied, sole cropping with Gliricidia sepium mulch applied, sole cropping with no mulch. The following parameters were measured: 1) plant-available soil P assessed by P uptake of maize and bean bioassay plants; 2) phosphate desorbable by anion exchange resin; 3) adsorption of added P into isotopically exchangeable and non-exchangeable pools.In the bioassay, P uptake of beans declined in the order: mulched sole-cropped>unmulched sole-cropped>alley-cropped soils. For maize the relative uptake was: mulched sole-cropped>unmulched sole-cropped = alley-cropped soils. These results suggest trees had not incorporated a significant quantity of P into the system after seven years and, probably, there was a decrease in available soil P due to the sequestration of P in the tree biomass. Potentially resin-desorbable P was higher in alley-cropped and mulched sole-cropped soils than in unmulched sole-cropped soils. The adsorption and desorption of added P into and from exchangeable and non-exchangeable pools did not differ between alley-cropped and unmulched sole-cropped soils.Crop yield and crop N, P and K uptake were all higher in the alley crops than in the unmulched sole crop. The supply of P to the crop under alley cropping seems to be dependent on P cycled and released from the mulch. The P cycle in alley cropping appears to be self-sustaining at least under conditions of moderate P fertiliser input.     

Plant rhizosphere influence on microbial C metabolism: the role of elevated CO2, N availability and root stoichiometry

Microbial decomposer C metabolism is considered a factor controlling soil C stability, a key regulator of global climate. The plant rhizosphere is now recognized as a crucial driver of soil C dynamics but specific mechanisms by which it can affect C processing are unclear. Climate change could affect microbial C metabolism via impacts on the plant rhizosphere. Using continuous ¹³C labelling under controlled conditions that allowed us to quantify SOM derived-C in all pools and fluxes, we evaluated the microbial metabolism of soil C in the rhizosphere of a C4 native grass exposed to elevated CO₂ and under variation in N concentrations in soil and in plant root C:N stoichiometry. Our results demonstrated that this plant can influence soil C metabolism and further, that elevated CO₂ conditions can alter this role by increasing microbial C efficiency as indicated by a reduction in soil-derived C respiration per unit of soil C-derived microbial biomass. Moreover, under elevated CO₂ increases in soil N, and notably, root tissue N concentration increased C efficiency, suggesting elevated CO₂ shifted the stoichiometric balance so N availability was a more critical factor regulating efficiency than under ambient conditions. The root C:N stoichiometry effect indicates that plant chemical traits such as root N concentration are able to influence the metabolism of soil C and that elevated CO₂ conditions can modulate this role. Increased efficiency in soil C use was associated with negative rhizosphere priming and we hypothesize that the widely observed phenomenon of rhizosphere priming may result, at least in part, from changes in the metabolic efficiency of microbial populations. Observed changes in the microbial community support that shifting microbial populations were a contributing factor to the observed metabolic responses. Our case study points at greater efficiency of the SOM-degrading populations in a high CO₂, high N world, potentially leading to greater C storage of microbially assimilated C in soil.     

Effect of mineral nitrogen on nitrogen nutrition and biomass partitioning between the shoot and roots of pea (Pisum sativum L.).

The effect of mineral N availability on nitrogen nutrition and biomass partitioning between shoot and roots of pea (Pisum sativum L., cv Baccara) was investigated under adequately watered conditions in the field, using five levels of fertiliser N application at sowing (0, 50, 100, 200 and 400 kg N ha^–1). Although the presence of mineral N in the soil stimulated vegetative growth, resulting in a higher biomass accumulation in shoots in the fertilised treatments, neither seed yield nor seed nitrogen concentration was affected by soil mineral N availability. Symbiotic nitrogen fixation was inhibited by mineral N in the soil but it was replaced by root mineral N absorption, which resulted in optimum nitrogen nutrition for all treatments. However, the excessive nitrogen and biomass accumulation in the shoot of the 400 kg N ha^–1 treatment caused crop lodging and slightly depressed seed yield and seed nitrogen content. Thus, the presumed higher carbon costs of symbiotic nitrogen fixation, as compared to root mineral N absorption, affected neither seed yield nor the nitrogen nutrition level. However, biomass partitioning within the nodulated roots was changed. The more symbiotic nitrogen fixation was inhibited, the more root growth was enhanced. Root biomass was greater when soil mineral N availability was increased: root growth was greater and began earlier for plants that received mineral N at sowing. Rooting density was also promoted by increased mineral N availability, leading to more numerous but finer roots for the fertilised treatments. However, the maximum rooting depth and the distribution of roots with depth were unchanged. This suggested an additional direct promoting effect of mineral N on root proliferation.