Detection of nitrogen transfer from N<Subscript>2</Subscript>-fixing shade trees to cacao saplings in <Superscript>15</Superscript>N labelled soil: ecological and experimental considerations

Theobroma cacao seedlings were grown alone (TCA) or associated with saplings of N₂-fixing shade trees Gliricidia sepium and Inga edulis in 200 l of ¹⁵N labelled soil within a physical root barrier for studying direct nitrogen transfer between the trees and cacao. Root:shoot partitioning ratio for sapling total N was lower than biomass root:shoot ratio in all species. Sapling total ¹⁵N was partitioned between root and shoot in about the same ratio as total N in cacao and inga but in gliricidia much higher proportion of ¹⁵N than total N was found in roots. Thus, whole plant harvesting should be used in ¹⁵N studies whenever possible. Average percentage of fixed N out of total tree N was 74 and 81% for inga estimated by a yield-independent and yield-dependent method, respectively, and 85% for gliricidia independently of estimation method. Strong isotopic evidence on direct N transfer from trees to cacao was observed in two cases out of ten with both tree species. Direct N transfer was not correlated with mycorrhizal colonisation of either donor or receiver plant roots. Direct N transfer from inga and gliricidia to cacao is conceivable but its prevalence and the transfer pathway via mycorrhizal connections or via reabsorption of N-rich legume root exudates by cacao require further study. Competition in the restricted soil space may also have limited the apparent transfer in this study because the trees accumulated more soil-derived N than cacao in spite of active N₂ fixation.     

Fallow enrichment of a traditional slash/mulch system in southern Costa Rica: comparisons of biomass production and crop yield

The slash and mulch system of frijol tapado or covered bean is a pre-hispanic system of bean production found throughout much of Central America. However, land use pressures have forced farmers to shorten the traditional fallow period, thus resulting in decreasing productivity. A potential solution is to enrich the fallow by using leguminous nitrogen-fixing trees. The enriched fallow systems evaluated in this study include both single and mixed species treatments: 1) Erythrina poeppigiana; 2) Calliandra colothrysus; 3) Gliricidia sepium; 4) Inga edulis; 5) Inga edulis and Erythrina poeppigiana; 6) Inga edulis and Calliandra calothrysus; and 7) Inga edulis and Gliricidia sepium. Biomass production of the fallow vegetation is shown to be of greater quality and quantity in all fallow enrichment treatments. However, bean yields did not show a significant response to the fallow enrichment treatments.     

Nitrogen contribution by multipurpose trees to rice and cowpea in an alley cropping system in Sierra Leone

In an alley cropping experiment, a study was carried out on N₂ fixation by Gliricidia sepium, nitrogen (N) accumulation by prunings of Gliricidia, Senna siamea (formerly Cassia siamea) and Gmelina arborea, and the N contribution to associated crops of rice and cowpea.Total N accumulated by the hedgerow trees ranged from 297–524 kg N ha^–1 on average but varied between tree species and depended on the growing season. Gliricidia sepium accumulated 370 kg N ha^–1 on average and more than half of this came from fixation. Senna siamea and Gmelina arborea served as reference trees for estimating N₂ fixation. The estimates of N₂ fixation using Gmelina as a reference gave higher estimates than those using Senna.Although the dry matter and nitrogen yields of prunings from the hedgerow trees were high, their relative nitrogen contribution to the associated crops was generally low ranging from 5 to 29%. Higher crop yields and nitrogen contribution were observed with Gliricidia sepium prunings. The low N contribution from prunings was attributed to the lack of synchronization between the N released from the prunings and the crop's demand for N.     

Cacao-coconut intercropping in Ghana: agronomic and economic perspectives

In Ghana, shade for cacao (Theobroma cacao L.) is becoming a critical issue because of extensive deforestation. Unlike in some other cacao-growing countries, cacao is not grown under the shade of coconut (Cocos nucifera) in Ghana. An experiment to compare the merits of four cacao-coconut intercropping systems with the traditional cultivation of cacao under Gliricidia sepium shade was undertaken at the Cocoa Research Institute of Ghana. Cacao seedling girth was not affected when intercropped with coconut but was significantly (P = 0.01) reduced when intercropped with G. sepium. High density cacao facilitated better early canopy formation. Yield of cacao spaced at 2.5 m triangular (1739 plants ha^–1) with coconut at 9.8 m triangular (105 plants ha^–1) was significantly higher (P = 0.05) than from the other treatments during 1993/94 to 1995/96. There were no major disease problems associated with intercropping cacao with coconuts. Widely spaced coconuts intercropped with cacao spaced at 3 m × 3 m showed better flowering and gave higher coconut yields, but cacao spaced at 2.5 m triangular under coconuts spaced at 9.8 m triangular was more profitable than the other treatments. Moisture stress was the greatest in cacao system with G. sepium shade and this could be responsible for the low yield of cacao in that treatment. It is suggested that properly arranged high density cacao under widely spaced coconuts can be a profitable intercrop system for adoption by cacao farmers in Ghana.This revised version was published online in November 2005 with corrections to the Cover Date.     

Measuring nitrogen fixation by <Emphasis Type="Italic">Sesbania sesban</Emphasis> planted fallows using <Superscript>15</Superscript>N tracer technique in Kenya

A field experiment was performed in eastern Kenya to estimate N₂ fixation by Sesbania sesban over an 18-month period using the ¹⁵N dilution method. The influence of three reference species, Senna spectabilis, Eucalyptus saligna and Grevillea robusta, on the estimates of N₂ fixation was also assessed. Percentage Ndfa (nitrogen derived from the atmosphere) was calculated based on foliar atom excess (FAE), above-ground atom excess (AAE) or whole tree atom excess (WAE) data. The differences in atom% ¹⁵N excess values between species and plant parts are presented and discussed. We recommend the use of several reference species for estimating %Ndfa and that the different results obtained should be carefully considered in relation to the issues being addressed. In this study, Senna was the most suitable of the three reference species because its N uptake pattern and phenology were very similar to those of Sesbania. When well established, the amount of N fixed by Sesbania accounts for more than 80% of its total N content, according to FAE-based estimates. We estimated the Ndfa by Sesbania after 18 months to between 500 and 600 kg ha⁻¹ , depending on whether FAE, AAE or WAE data were used and on the choice of reference species. The substantial accumulation of N in planted Sesbania highlighted its potential to increase the sustainability of crop production on N-limited soils. We consider the ¹⁵N dilution method to be appropriate for quantifying N₂ fixation in improved fallows in studies, similar to this one, of young trees with high N₂-fixing ability.     

Nitrogen-fixation dynamics in a cut-and-carry silvopastoral system in the subhumid conditions of Guadeloupe, French Antilles

This paper summarizes several studies on N recycling in a tropical silvopastoral system for assessing the ability of the system to increase soil fertility and insure sustainability. We analyzed the N₂ fixation pattern of the woody legume component (Gliricidia sepium), estimated the recycling rate of the fixed N in the soil, and measured N outputs in tree pruning and cut grass (Dichanthium aristatum). With this information, we estimated the N balance of the silvopastoral system at the plot scale. The studies were conducted in an 11-year-old silvopastoral plot established by planting G. sepium cuttings at 0.3 m × 2 m spacing in natural grassland. The plot was managed as a cut-and-carry system where all the tree pruning residues (every 2-4 months) and cut grass (every 40-50 days) were removed and animals were excluded. No N fertilizer was applied. Dinitrogen fixation, as estimated by the ¹⁵N natural abundance method, ranged from 60-90% of the total N in aboveground tree biomass depending on season. On average, 76% of the N exports from the plot in tree pruning (194 kg [N] ha^–1 yr^–1) originated from N₂ fixation. Grass production averaged 13 Mg ha^–1 yr^–1 and N export in cut grass was 195 kg [N] ha^–1 yr^–1. The total N fixed by G. sepium, as estimated from the tree and grass N exports and the increase in soil N content, was about 555 kg [N] ha^–1 yr^–1. Carbon sequestration averaged 1.9 Mg [C] ha^–1 yr^–1 and soil organic N in the 0-0.2 m layer increased at a rate of 166 kg [N] ha^–1 yr^–1, corresponding to 30% of N₂ fixation by the tree. Nitrogen released in nodule turnover (10 kg [N] ha^–1 yr^–1) and litter decomposition (40 kg [N] ha^–1 yr^–1) contributed slightly to this increase, and most of the recycled N came from the turnover or the activity of other below-ground tree biomass than nodules. This revised version was published online in June 2006 with corrections to the Cover Date.     

Mixed-species plantations of Acacia mangium and Eucalyptus grandis in Brazil: 2: Nitrogen accumulation in the stands and biological N2 fixation

The sustainability of plantation forests is closely dependent on soil nitrogen availability in short-rotation forests established on low-fertility soils. Planting an understorey of nitrogen-fixing trees might be an attractive option for maintaining the N fertility of soils. The development of mono-specific stands of Acacia mangium (100A:0E) and Eucalyptus grandis (0A:100E) was compared with mixed-species plantations, where A. mangium was planted in a mixture at a density of 50% of that of E. grandis (50A:100E). N₂ fixation by A. mangium was quantified in 100A:0E and 50A:100E at age 18 and 30 months by the ¹⁵N natural abundance method and in 50A:100E at age 30 months by the ¹⁵N dilution method. The consistency of results obtained by isotopic methods was checked against observations of nodulation, Specific Acetylene Reduction Activity (SARA), as well as the dynamics of N accumulation within both species. The different tree components (leaves, branches, stems, stumps, coarse roots, medium-sized roots and fine roots) were sampled on 5–10 trees per species for each age. Litter fall was assessed up to 30 months after planting and used to estimate fine root mortality. Higher N concentrations in A. mangium tree components than in E. grandis might be a result of N₂ fixation. However, no evidence of N transfer from A. mangium to E. grandis was found. SARA values were not significantly different in 100A:0E and 50A:100E but the biomass of nodules was 20–30 times higher in 100A:0E than in 50A:100E. At age 18 months, higher δ¹⁵N values found in A. mangium tree components than in E. grandis components prevented reliable estimations of the percentage of N derived from atmospheric fixation (%Ndfa). At age 30 months, %Ndfa estimated by natural abundance and by ¹⁵N dilution amounted to 10–20 and 60%, respectively. The amount of N derived from N₂ fixation in the standing biomass was estimated at 62 kg N ha⁻¹ in 100A:0E and 3 kg N ha⁻¹ in 50A:100E by the ¹⁵N natural abundance method, and 16 kg N ha⁻¹ in 50A:100E by the ¹⁵N dilution method. The total amount of atmospheric N₂ fixed since planting (including fine root mortality and litter fall) was estimated at 66 kg N ha⁻¹ in 100A:0E and 7 kg N ha⁻¹ in 50A:100E by the ¹⁵N natural abundance method, and 31 kg N ha⁻¹ in 50A:100E by the ¹⁵N dilution method. The most reliable estimation of N₂ fixation was likely to be achieved using the ¹⁵N dilution method and sampling the whole plant.     

Testing the safety-net role of hedgerow tree roots by 15N placement at different soil depths

Trees which root below crops may have a beneficial role in simultaneous agroforestry systems by intercepting and recycling nutrients which leach below the crop rooting zone. They may also compete less strongly for nutrients than trees which root mainly within the same zone as crops. To test these hypotheses we placed highly enriched ¹⁵N-labelled ammonium sulphate at three depths in the soil between mixed hedgerows of the shallow-rooting Gliricidia sepium and the deep rooting Peltophorum dasyrrhachis. A year after the isotope application most of the residual ¹⁵N in the soil remained close to the injection points due to the joint application with a carbon source which promoted ¹⁵N immobilization. Temporal ¹⁵N uptake patterns (two-weekly leaf sub-sampling) as well as total ¹⁵N recovery measurements suggested that Peltophorum obtained more N from the subsoil than Gliricidia. Despite this Gliricidia appeared to compete weakly with the crop for N as it recovered little ¹⁵N from any depth but obtained an estimated 44–58% of its N from atmospheric N₂-fixation. Gliricidia took up an estimated 21 kg N ha^–1 and Peltophorum an estimated 42 kg N ha^–1 from beneath the main crop rooting zone. The results demonstrate that direct placement of ¹⁵N can be used to identify N sourcing by trees and crops in simultaneous agroforestry systems, although the heterogeneity of tree root distributions needs to be taken into account when designing experiments. This revised version was published online in June 2006 with corrections to the Cover Date.     

Crop residue effect on crop performance,soil N<Subscript>2</Subscript>O and CO<Subscript>2</Subscript> emissions in alley cropping systems in subtropical China

Land management practices that simultaneously improve soil properties are crucial to high crop production and minimize detrimental impact on the environment. We examined the effects of crop residues on crop performance, the fluxes of soil N₂O and CO₂ under wheat-maize (WM) and/or faba bean-maize (FM) rotations in Amorpha fruticosa (A) and Vetiveria zizanioides (V) intercropping systems on a loamy clay soil, in subtropical China. Crop performance, soil N₂O and CO₂ as well as some potential factors such as soil water content, soil carbon, soil nitrogen, microbial biomass and N mineralization were recorded during 2006 maize crop cultivation. Soil N₂O and CO₂ fluxes are determined using a closed-based chamber. Maize yield was greater after faba bean than after wheat may be due to differences in supply of N from residues. The presence of hedgerow significantly improved maize grain yields. N₂O emissions from soils with maize were considerably greater after faba bean (345 g N₂O–N ha⁻¹) than after wheat (289 g N₂O–N ha⁻¹). However, the cumulated N₂O emissions did not differ significantly between WM and FM. The difference in N₂O emissions between WM and FM was mostly due to the amounts of crop residues. Hedgerow alley cropping tended to emit more N₂O than WM and FM, in particular A. fruticosa intercropping systems. Over the entire 118 days of measurement, the N₂O fluxes represented 534 g N₂O–N ha⁻¹ (AWM) and 512 g N₂O–N ha⁻¹ (AFM) under A. fruticosa species, 403 g N₂O–N ha⁻¹ (VWM) and 423 g N₂O–N ha⁻¹ (VFM) under Vetiver grass. We observed significantly higher CO₂ emission in AFM (5,335 kg CO₂–C ha⁻¹) from June to October, whereas no significant difference was observed among WM (3,480 kg CO₂–C ha⁻¹), FM (3,302 kg CO₂–C ha⁻¹), AWM (3,877 kg CO₂–C ha⁻¹), VWM (3,124 kg CO₂–C ha⁻¹) and VFM (3,309 kg CO₂–C ha⁻¹), indicating the importance of A. fruticosa along with faba bean residue on CO₂ fluxes. As a result, crop residues and land conversion from agricultural to agroforestry can, in turn, influence microbial biomass, N mineralization, soil C and N content, which can further alter the magnitude of crop growth, soil N₂O and CO₂ emissions in the present environmental conditions.     

Positive nitrogen balance of <Emphasis Type="Italic">Acacia mangium</Emphasis> woodlots as fallows in the Philippines based on <Superscript>15</Superscript>N natural abundance data of N<Subscript>2</Subscript> fixation

Nitrogen inputs from biological nitrogen fixation contribute to productivity and sustainability of agroforestry systems but they need to be able to offset export of N when trees are harvested. This study assessed magnitudes of biological nitrogen fixation (natural ¹⁵N abundance) and N balance of Acacia mangium woodlots grown in farmer’s fields, and determined if N₂ fixation capacity was affected by tree age. Tree biomass, standing litter, understory vegetation and soil samplings were conducted in 15 farmer’s fields growing A. mangium as a form of sequential agroforestry in Claveria, Misamis Oriental, Philippines. The trees corresponded to ages of 4, 6, 8, 10 and 12 years, and were replicated three times. Samples from different plant parts and soils (0–100 cm) were collected and analyzed for δ¹⁵N and nutrients. The B-value, needed as a reference of isotopic discrimination when fully reliant on atmospheric N, was generated by growing A. mangium in an N₂-free sand culture in the glasshouse. Isotopic discrimination occurring during N₂ fixation and metabolic processes indicated variation of δ¹⁵N values in the order of nodules > old leaves > young leaves > stems > litterfall and roots of the trees grown in the field, with values ranging from −0.8 to 3.5‰ except nodules which were enriched and significantly different from other plant parts (P < 0.0001). Isotopic discrimination was not affected by tree age (P > 0.05). Plants grown in N free sand culture exhibited the same pattern of isotopic discrimination as plants grown in the field. The estimated B-value for the whole plant of A. mangium was −0.86‰. Mature tree stands of 12 years accumulated up to 1994 kg N ha⁻¹ in aboveground biomass. Average proportion of N derived from N₂ fixation of A. mangium was 54% (±22) and was not affected by age (P > 0.05). Average yearly quantities of N₂ fixed were 128 kg N ha⁻¹ in above-ground biomass amounting to 1208 kg N fixed ha⁻¹ over 12 years. Harvest of 12-year old trees removed approximately 91% of standing aboveground biomass from the site as timber and fuel wood. The resulting net N balance was +151 kg N ha⁻¹ derived from remaining leaves, twigs, standing litter, and +562 kg N ha⁻¹ when tree roots were included in the calculation. The fast growing A. mangium appears to be a viable fallow option for managing N in these systems. However, other nutrients have to be replaced by using part of the timber and fuel wood sales to compensate for large amounts of nutrient removed in order for the system to be sustainable.     

Response of <Emphasis Type="Italic">Theobroma cacao</Emphasis> and <Emphasis Type="Italic">Inga edulis</Emphasis> seedlings to cross-inoculated populations of arbuscular mycorrhizal fungi

Response of Theobroma cacao L. (cacao) and the shade tree Inga edulis Mart. (inga) seedlings from an organically-grown cacao plantation to inoculation with native arbuscular mycorrhizae forming fungi (AMF) was studied in a cross-inoculation assay under greenhouse conditions. Seedlings of inga and cacao were grown in pots filled with heat-treated soil from the plantation. Control was heat-treated soil without inoculum and roots of cacao and inga from the plantation were applied as AMF inocula. Undisturbed soil blocks were used as a “positive control” of the inoculation potential of untreated soil and roots combined. No AMF structures were observed in the roots of either species in the heat-treated control. All inocula were infective in both hosts and the differences in the total AMF colonization percentage between the hosts were not significant but inga had significantly higher colonization by hyphal coils and arbuscules. Cacao roots but neither inga roots nor soil block inocula stimulated cacao growth. All inocula significantly increased growth of inga, which had higher relative mycorrhizal responsiveness than cacao. Thus, in spite of the strong infectivity of the inocula in both hosts, cacao and inga responded differently to the same AMF populations. The strong conspecific preference of cacao suggests that attention must be paid to the AMF inoculum used for this species. However, the strong response of inga to cacao root inoculum indicates that the two species may share same AMF symbionts, thus enabling positive interactions between them, including formation of common mycelial networks.     

Do multipurpose companion trees affect high value timber trees in a silvopastoral plantation system?

Establishment of native timber trees on deforested land may contribute to the livelihood of farmers, to improved ecosystem services and to increased greenhouse gas uptake. Here, we present a new silvopastoral planting design to assess species performance and interspecific competition or facilitation effects among native timber and multipurpose trees in Central America. Two timber species, Tabebuia rosea and Cedrela odorata, were established in three low-density planting regimes allowing combined tree and future livestock production: (1) solitary planting, (2) companion planting with Guazuma ulmifolia, and (3) companion planting with the nitrogen-fixing Gliricidia sepium. We quantified survival, growth and reforestation potential of the two timber species subjected to the different planting regimes for the first 2 years after establishment. Nitrogen concentration as well as stable nitrogen and carbon isotope composition (δ¹⁵N, δ¹³C) of leaves of the timber saplings were determined. T. rosea showed higher survival and better growth than C. odorata under varying environmental conditions (soil, concomitant vegetation). Performance of the timber saplings was unaffected by either companion species. Planting regimes had no effect on foliar nitrogen concentration and δ¹⁵N of the two timber species, although δ¹⁵N values indicated nitrogen fixation activity in G. sepium trees. Planting regimes affected foliar δ¹³C values in T. rosea. δ¹³C values were significantly higher in solitarily growing individuals, suggesting lower exposition to water stress conditions in saplings surrounded by companion species. As we found positively correlated growth traits among timber and multipurpose trees, a combined planting may benefit farmers by providing additional goods and services.     

Foliar Carbon Isotope Composition (δ 13C) and Water Use Efficiency of Different Populus deltoids Clones Under Water Stress

Foliar carbon isotope composition (δ ¹³C), total dry biomass, and long-term water use efficiency (WUE_L) of 12 Populus deltoids clones were studied under water stress in a greenhouse. Total dry biomass of clones decreased greatly, while δ ¹³C increased. Single-element variance analysis in the same water treatment indicated that WUE_L difference among clones was significant. Clones J₂, J₆, J₇, J₈, and J₉ were excellent with high WUE_L. Extremely significant δ ¹³C differences among water treatments and clones were revealed by two-element variance analysis. Water proved to be the primary factor affecting δ ¹³C under water stress. It showed that there was a good positive correlation between δ ¹³C and WUE_L in the same water treatment, and that a high WUE_L always coincided with a high δ ¹³C. δ ¹³C might be a reliable indirect index to estimate WUE_L among P. deltoids clones. Translated from Scientia Silvae Sinicae, 2005, 41(1) (in Chinese)     

Anthropogenic disturbance of natural forest vegetation on calcareous soils alters soil organic matter composition and natural abundance of <Superscript>13</Superscript>C and <Superscript>15</Superscript>N in density fractions

In the last century, many calcareous soils in Castilla León (northwestern Spain) have been transformed from natural Quercus ilex forest to cropped land. Reforestation with Pinus halepensis has been taking place during the past 40 years. In order to obtain a better understanding of how these disturbances affect ecosystem functioning, we studied the quantity and quality of soil organic matter (SOM) in natural forest ecosystems, cropland and Pinus plantations. Density fractionation combined with ultrasonic dispersion enables separation and study of SOM fractions: free organic matter (OM), OM occluded into aggregates and OM stabilized in organo-mineral complexes, considered on the basis of the type of physical protection provided. We separated SOM density fractions and determined the concentrations of C and N, C/N ratios and the natural isotopic abundance (δ¹³C and δ¹⁵N values). Transformation of Quercus forest to cropland resulted in major losses of SOC and N, as expected. However, subsequent reforestation with Pinus resulted in good recovery of the original SOC and soil N pools. This indicates the potential for enhanced C storage in agricultural soils by their reversion to a forested state. Study of the density fractions and their ¹³C and ¹⁵N signatures enabled better understanding of the high stability of OM in calcareous soils, and analysis of δ¹³C variations throughout the profile also enabled identification of past C3/C4 vegetation change. Despite the different OC contents of soils under different land use, OM stabilization mechanisms were not significantly different. In calcareous soils, accumulation of SOC and N is mainly due to organo-mineral associations, resulting in physicochemical stabilization against further decomposition.     

Contributions of salmon-derived nitrogen to riparian vegetation in the northwest Pacific region

We examined the relationship between the annual escapement of salmon and the δ ¹⁵N of willow (Salix spp.) leaves to evaluate the contribution of marine-derived nutrients (MDN) to riparian vegetation around the Pacific Northwest and Northeast regions. Foliar δ ¹⁵N values ranged from −3.42‰ to 4.65‰. The value increased with increasing density of carcasses up to 500 fish/km and 1500 fish/km. δ ¹⁵N values were variable at carcass densities below 500 fish/km. Possible factors affecting the fluctuation of δ ¹⁵N at reference sites are: (1) denitrification; (2) the presence of N₂-fixing trees, such as alder; and (3) agricultural runoff. δ ¹⁵N values at the sites with carcass densities over 500 fish/km were consistently high, while a value of δ ¹⁵N below zero was observed at only one site (Rusha River; δ ¹⁵N = −1.87‰). At this site, most adult pink salmon returned to limited locations near the estuary because steeper channel gradients acted as a migration barrier, resulting in the negative δ ¹⁵N value. Nevertheless, we concluded that our results showed evidence of the feedback of MDN to terrestrial vegetation, although the use of the δ ¹⁵N value as a terrestrial end member at spawning sites is limited. If the relationship between the enrichment index, which is expressed as the values using a mixing model, and salmon abundance was estimated, the availability of MDN in riparian ecosystems could possibly be evaluated and will lead to the establishment of escapement goals. An erratum to this article is available at .     

Nutrient contents and efficiencies of beech and spruce saplings as influenced by competition and O<Subscript>3</Subscript>/CO<Subscript>2</Subscript> regime

Saplings of Fagus sylvatica and Picea abies were grown under conditions of intra and interspecific competition in a 2-year phytotron study under combinations of ambient and elevated ozone (+O₃ which is 2 × O₃, but <150 nl l⁻¹) as well as carbon dioxide concentrations (+CO₂ which is amb. CO₂ + 300 μl CO₂ l⁻¹) in a full factorial design. Saplings were analysed for various mineral nutrients in different plant organs as well as biomass production and crown development. The study was based on the assumption that nutritional parameters important for growth and competitiveness are affected by stress defence under limiting nutrient supply. The hypotheses tested were (1) that nutrient uptake-related parameters (a) as well as efficiencies in nutrient use for above-ground competition (b) of beech rather than spruce are impaired by the exposure to elevated O₃ concentrations, (2) that the efficiency in nutrient uptake of spruce is enhanced by elevated CO₂ concentrations in mixed culture, and (3) that the ability to occupy above-ground space at low nutrient cost is co-determinant for the competitive success in mixed culture. Clear nitrogen deficiencies were indicated for both species during the 2-year phytotron study, although foliar nitrogen-biomass relationships were not so close for spruce than for beech. O₃ stress did not impair nutrient uptake-related parameters of beech; thus hypothesis (1a). was not supported. A negative effect of elevated O₃ (under amb. CO₂) on the N and P based efficiencies in above-ground space occupation (i.e. lower crown volume per unit of N or P invested in stems, limbs and foliage) of beech supported hypothesis (1b). It appeared that ozone stress triggered a nutrient demand for stress defence and tolerance at the expense of above-ground competition (trade-off). Crown volume of beech under O₃ stress was stabilized in monoculture by increased nutrient uptake. In general, the +CO₂-treatment was able to counteract the impacts of 2 × O₃. Elevated CO₂ caused lower N and S concentrations in current-year foliage of both tree species, slightly higher macronutrient amounts in the root biomass of spruce, but did not increase the efficiencies in nutrient uptake of spruce in mixed culture. Therefore hypothesis (2) was not supported. At the end of the experiment spruce turned out to be the stronger competitor in mixed culture as displayed by its higher total shoot biomass and crown volume. The amounts of macronutrients in the above-ground biomass of spruce individuals in mixed culture distinctly exceeded those of beech, which had been strongly reduced by interspecific competition. The superior competitiveness of spruce was related to higher N and P-based efficiencies in above-ground space occupation as suggested in hypothesis (3). This article belongs to the special issue “Growth and defence of Norway spruce and European beech in pure and mixed stands”.     

Vesicular-arbuscular mycorrhiza effects on Gliricidia sepium and Senna siamea in a fallowed alley cropping system

Nitrogen fixing and non-N₂ fixing legumes such as Gliricidia speium and Senna siamea have been used in alley cropping systems for soil improvement and source of N for food crops. However their establishments could be limited by P and moisture deficiencies in degraded soils. Vesicular-arbuscular mycorrhizal fungi can help to overcome these deficiencies. We examined the effects of a vesicular-arbuscular mycorrhizal (VAM) fungus, Glomus deserticola, on the performance of sole hedgerow trees of Gliricidia sepium and Senna siamea and their mixtures (interplanted) in a fallowed alley cropping experiment on a degraded Alfisol in southwestern Nigeria. Percentage root infection by VAM fungi was higher in inoculated plants than in uninoculated ones irrespective of whether they were interplanted or non-interplanted. Inoculation with G. deserticola increased dry matter accumulation and nutrient uptake (N. P, Mg and K) but there was no significant interaction between mycorrhizal inoculation and interplanting for growth and nutrient uptake except for the uptake of P, Mg and K in G. sepium. Inoculation with G. deserticola reduced leaf shedding of G. sepium by 50% but did not have the same effect for S. siamea. For both tree species inoculated plants extracted more water from 0–30 cm depth than the uninoculated ones.     

Mixed planted-fallows using coppicing and non-coppicing tree species for degraded Acrisols in eastern Zambia

The widespread planting of Sesbania sesban fallows for replenishing soil fertility in eastern Zambia has the potential of causing pest outbreaks in the future. The pure S. sesban fallows may not produce enough biomass needed for replenishing soil fertility in degraded soils. Therefore, an experiment was conducted at Kagoro in Katete district in the Eastern Province of Zambia from 1997 to 2002 to test whether multi-species fallows, combining non-coppicing with coppicing tree species, are better than mono-species fallows of either species for soil improvement and increasing subsequent maize yields. Mono-species fallows of S. sesban (non-coppicing), Gliricidia sepium, Leucaena leucocephala and Acacia angustissima (all three coppicing), and mixed fallows of G. sepium + S. sesban, L. leucocephala + S. sesban, A. angustissima + S. sesban and natural fallow were compared over a three-year period. Two maize (Zea mays) crops were grown subsequent to the fallows. The results established that S. sesban is poorly adapted and G. sepiumis superior to other species for degraded soils. At the end of three years, sole G. sepium fallow produced the greatest total biomass of 22.1 Mg ha⁻¹ and added 27 kg ha⁻¹ more N to soil than G. sepium + S. sesban mixture. During the first post-fallow year, the mixed fallow at 3.8 Mg ha⁻¹ produced 77% more coppice biomass than sole G. sepium, whereas in the second year both sole G. sepium and the mixture produced similar amounts of biomass (1.6 to 1.8 Mg ha⁻¹). The G. sepium + S. sesban mixture increased water infiltration rate more than sole G. sepium, but both these systems had similar effects in reducing soil resistance to penetration compared with continuous maize without fertilizer. Although sole G. sepium produced high biomass, it was G. sepium + S. sesban mixed fallow which resulted in 33% greater maize yield in the first post-fallow maize. However, both these G. sepium-based fallows had similar effects on the second post-fallow maize. Thus the results are not conclusive on the beneficial effects of G. sepium + S. sesban mixture over sole G. sepium. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of the application of the leaf mulch of Gliricidia sepium on early development,leaf nutrient contents and tuber yields of water yam (Dioscorea alata)

This paper describes the effects of the leaf mulch of Gliricidia sepium on the development and yield of Water Yam, Dioscorea alata. Using leaf mulch, the time taken for the yam setts to sprout can be shortened by approximately 20 percent.Organic mulches contain considerable quantities of plant nutrients. Increasing amounts of mulch improved the leaf nutrient contents of the yam crop and resulted in significantly higher tuber yields. Over a tuber yield range up to c. 15 tons ha^–1 each additional ton DM Gliricidia sepium mulch applied resulted in a yield increment of about 2 ton yam tubers. A nutrient supply — nutrient extraction balance is discussed, comparing mulch applied and yam tubers harvested.Mulching as agricultural technique is a useful and affordable tool in adapting low external input cropping systems to local economic and environmental conditions.     

Soil nitrogen as affected by Gliricidia sepium in a silvopastoral system in Guadeloupe, French Antilles

Knowledge of the status and dynamics of soil N is essential to improving the production and management of silvopastoral system in the tropics. Soil N status and dynamics were analyzed as key factors affecting productivity and sustainability of a cut-and-carry silvopastoral system. The total soil N and N mineralization as affected by soil moisture and temperature were studied in a Gliricidia sepium (Jacq.) Walp – Dichanthium aristatum (Poir) C.E. Hubbard grassland association and in an adjacent open grassland located in a subhumid tropical region. The plot was installed in 1989 and the pruning residues and cut grass were removed from the site. No N fertilizer was applied. Total soil N increased at an average rate of 180 kg N ha⁻¹ in the 0–0.2 m soil layer of the silvopastoral plot. Only a third of this value could be explained by the litter, nodule and root turnover. Nitrogen mineralization in both soils varied as a function of temperature but it was not affected by soil moisture. Even if mean soil temperature was 1 °C to 2 °C greater in the open grassland, the estimated daily rate of in-situ N mineralization was 20% greater in the silvopastoral system. Our results indicated that greater N mineralization in the silvopastoral system was due to greater soil biological activity associated with higher soil organic matter rather than due to more favorable soil temperature and water conditions. This revised version was published online in June 2006 with corrections to the Cover Date.