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.     

Nitrogen contribution of five leguminous trees and shrubs to alley cropped maize in Ibadan, Nigeria

There are abundant local legume trees and shrubs potentially suitable for alley cropping systems in the sub-Saharan Africa, which are yet to be studied. The nitrogen contribution of two years old Albizia lebbeck and S. corymbosato yield of maize grown in alley cropping was compared to that of Senna siamea, Gliricidia sepium and Leucaena leucocephala in four seasons at Ibadan. Maize shoot biomass and maize grain yield in A. lebbeck alley compared favourably with that in G. sepium and L. leucocephala. Maize biomass and grain yield in S. corymbosa alleys were the lowest. Within A. lebbeck, L. leucocpehala, and G. sepium alleys there were no significant differences in the maize yield in the alleys that received 0, 40 or 80 kg N/ha. Application of more than 40 kg N/ha in S. corymbosa alleys was not necessary as there was no significant increase in maize yield at the higher level of nitrogen. Maize yield and N uptake in A. lebbeck alleys were not significantly different from yield and N uptake in G. sepium, and L. leucocephala at the same fertilizer level. There was a significant correlation between hedgerow tree biomass and maize grain yield. At the end of twelve weeks after pruning application, the organic residues of the pruning applied in the alleys ranged from 5% in G. sepium and 44% in A. lebbeck in the first year compared with the original pruning applied which showed that the slow rate of A. lebbeck decomposition could have a beneficial effect on the soil. The maize N recovery from applied N fertilizer was low (10–22%). Percentage N recovery from the prunings was low in the non-N fixing trees (12–22%), while the recovery was high (49–59%) in A. lebbeck as well as in the other nitrogen fixing tree prunings. Thus A. lebbeck, apart from enhancing maize growth and grain yield like in L. leucocephala and G. sepium, had an added advantage because it remained longer as mulching material on the soil because of its slow rate of decomposition. It was able to survive pruning frequencies with no die-back. This indicates that A. lebbeck is a good potential candidate for alley cropping system in West Africa. S. corymbosa performed poorly compared with the other legume trees. Though it responded to N fertilizer showing a positive interaction between the hedgerow and fertilizer application, it had a high die back rate following pruning periods and termite attack.This revised version was published online in November 2005 with corrections to the Cover Date.     

Selective control of weeds in an arable crop by mulches from some multipurpose trees in Southwestern Nigeria

The use of agroforestry systems in which pruning from trees is used to mulch the companion crops is an important area of research in the tropics. However, previous studies mostly evaluated the contribution of mulch to soil improvement and rarely examined the effect of mulch on weeds. Field experiments were conducted during the 1995 and 1996 growing seasons to investigate the effects of mulch from three woody fallow species on weed composition, biomass and maize grain yield. Treatments consisted of mulch from Leucaena leucocephala, Gliricidia sepium, and Senna siamea applied at rates of five and three tons dry matter ha^–1 at planting and three weeks after planting (WAP), respectively, an unmulched treatment that received 90 kg N ha^–1 of inorganic fertiliser, and an unmulched control plot that received no fertiliser. In both years and sampling dates, plots mulched with G. sepium and S. siamea had significantly lower weed density and biomass than the control plot in each of the sampling times and year of study. There was no significant difference in either weed density or biomass between the plot mulched with L. leucocephala and the unmulched plots. Mulches from G. sepium and S. siamea reduced weed density and weed biomass, while L. leucocephala was less effective in reducing weed biomass and weed density. Weed reduction by the mulches was in the order G. sepium S. siamea > L. leucocephala. Sedges were the dominant species in all the treatments except in G. sepium plots, where Talinum triangulare and other broadleaved species were dominant.This revised version was published online in November 2005 with corrections to the Cover Date.     

Effect of tree pruning and pruning application to trees on nitrogen fixation by Leucaena and Gliricidia

A microplot study was conducted with Leucaena leucocephala and Gliricidia sepium to determine the effect of pruning the trees and applying the prunings to the soil (as either surface mulch or soil-incorporated green manure) on N₂ fixation by the trees. N₂ fixation was estimated by the ¹⁵N dilution method, and the non-N₂-fixing species Senna siamea was used as a reference species. Number and dry weight of nodules were reduced with applications of prunings in Leucaena, and increased in Gliricidia. For both species, the lowest rate of N₂ fixation (47%) occurred under green manure incorporation after pruning, and the highest (66%) with mulching and no-pruning. Incorporation of green manure into soil significantly depressed N₂ fixation by 19.1% in Gliricidia and 20.6% in Leucaena compared to no-pruning treatment. The deleterious effect of incorporation of prunings into soil also affected the amount of N₂ fixed. During the nine months following pruning, green manure incorporation steadily decreased the percentage of N₂ fixed, whereas with no-pruning, pruning only and pruning with mulching, N₂ fixation increased with time. Reduction of N₂ fixation in pruned plants could imply that they would compete for N with interplanted food crops, causing lower N use efficiency by the latter. Despite lower dry-matter and N yields of Leucaena as compared with Gliricidia, its biological N₂ fixation amounted to 73% of its N requirement as opposed to the corresponding figures of 50% for Gliricidia. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

Natural abundance of <Superscript>15</Superscript>N in two cacao plantations with legume and non-legume shade trees

Natural abundance of ¹⁵N was sampled in young and mature leaves, branches, stem, and coarse roots of trees in a cacao (Theobroma cacao) plantation shaded by legume tree Inga edulis and scattered non-legumes, in a cacao plantation with mixed-species shade (legume Gliricidia sepium and several non-legumes), and in a tree hedgerow bordering the plantations in Guácimo, in the humid Caribbean lowlands of Costa Rica. The deviation of the sample ¹⁵N proportion from that of atmosphere (δ¹⁵N) was similar in non-legumes Cordia alliodora, Posoqueria latifolia, Rollinia pittieri, and T. cacao. Deep-rooted Hieronyma alchorneoides had lower δ¹⁵N than other non-N₂-fixers, which probably reflected uptake from a partially different soil N pool. Gliricidia sepium had low δ¹⁵N. Inga edulis had high δ¹⁵N in leaves and branches but low in stem and coarse roots. The percentage of N fixed from atmosphere out of total tree N (%N_f) in G. sepium varied 56–74%; N₂ fixation was more active in July (the rainiest season) than in March (the relatively dry season). The variation of δ¹⁵N between organs in I. edulis was probably associated to ¹⁵N fractionation in leaves. Stem and coarse root δ¹⁵N was assumed to reflect the actual ratio of N₂ fixation to soil N uptake; stem-based estimates of %N_f in I. edulis were 48–63%. Theobroma cacao below I. edulis had lower δ¹⁵N than T. cacao below mixed-species shade, which may indicate direct N transfer from I. edulis to T. cacao but results so far were inconclusive. Further research should address the ¹⁵N fractionation in the studied species for improving the accuracy of the N transfer estimates. The δ¹⁵N appeared to vary according to ecophysiological characteristics of the trees.     

Soil property changes in contour hedgerow systems on sloping land in the Philippines

The impact of contour hedgerow systems on soil sustainability under acidic conditions has been widely criticized. A study was undertaken to determine the effects of management and hedgerow species on soil properties. Cassia spectabilis (a non-N-fixing tree legume), Gliricidia sepium (an N-fixing tree legume), Pennisetum purpureum (a forage grass), and Stylosanthes guyanensis (a forage legume) contour barriers were compared with an open field (non-hedgerow treatment) over 1 cowpea and 2 rice seasons. Three types of management viz.: prunings applied + N₀P₀K₀, prunings applied + N₅₀P₂₀K₂₀ and prunings removed + N₅₀P₂₀K₂₀ were used as subplot treatments. The soils were strongly acidic (pH 4.5) and classified as clay Orthoxic Palehumult. Cassia performed better than the other species in terms of pruning biomass, N and P contributions over a period of 20 months. There was a combined positive effect of pruning biomass and fertilizers on rice and cowpea yields in Pennisetum and Gliricidia systems, while a tendency towards a positive effect of pruning biomass on rice was found in the Cassia system. The pruning biomass and/or fertilizer application did not significantly influence the top soil organic C, N and available P in the hedgerow systems. Soil bulk density was significantly reduced by the application of Cassia prunings after 12 months. Organic C, N and P dynamics indicated that in situ pruning biomass was not sufficient to maintain their level in the soil. But the cassia systems with prunings applied + N₅₀P₂₀K₂₀ experienced the lowest degradation in soil organic C (2.1 t ha⁻¹) followed by the Gliricidia systems (4.1 t ha⁻¹). The overall results imply that the application of pruning and inorganic fertilizer is imperative to conserve soil resources, and non- N-fixing tree species can exert a significant advantage in biomass and thereby in soil N-recycling under acidic soil. This revised version was published online in June 2006 with corrections to the Cover Date.     

Decreasing rice and cowpea yields in alley cropping on a highly weathered Oxisol in West Sumatra,Indonesia

A hedgerow intercropping study was conducted for 7 years in West Sumatra, Indonesia on an acid and highly A1-saturated (72%) soil to determine growth and yield responses of tree hedgerows and upland rice and cowpea intercrops. Three tree species,Paraserianthes falcataria, Calliandra calothyrsus, andGliricidia sepium, and a no tree control were planted at three lime rates of zero, 375 kg ha^–1, and liming to 25% Al+H saturation. Annual fertilizer inputs of 20 kg P and 50 kg K ha^–1 were kept low to approximate low input farming systems. The trees were pruned 4 to 6 times per year and prunings applied to the intercrops.Paraserianthes andCalliandra grew vigorously, whileGiricidia grew poorly and was replaced after four years withFlemingia macrophylla. Hedgerow growth and yields were reported in a previous paper [Evensen et al., 1994].Rice and cowpea yields initially increased with lime andParaserianthes pruning application, but yields and soil cations (Ca, Mg, and K) declined until fertilizer inputs were increased after four years. Thereafter, crop yields increased and soil cations returned to original levels. Soil C and N were maintained over the 7 years on plots with trees. These results indicate little build-up of nutrient cations due to recycling by the trees and suggest that successful alley farming on such highly weathered soils requires maintenance of soil fertility with external inputs.     

Above- and below-ground biomass dynamics in a sole cropping and an alley cropping system withGliricidia sepium in the semi-deciduous rainforest zone of West Africa

A considerable amount of data is available about above-ground biomass production and turnover in tropical agroforestry systems, but quantitative information concerning root turnover is lacking. Above- and below-ground biomass dynamics were studied during one year in an alley cropping system withGliricidia sepium and a sole cropping system, on aPlinthic Lixisol in the semi-deciduous rainforest zone of the Côte d'Ivoire. Field crops were maize and groundnut. Live root mass was higher in agroforestry than in sole cropping during most of the study period. This was partly due to increased crop and weed root development and partly to the presence of the hedgerow roots. Fine root production was higher in the alleys and lower under the hedgerows compared to the sole cropping plots. Considering the whole plot area, root production in agroforestry and sole cropping systems was approximatly similar with 1000–1100 kg ha^–1 (dry matter with 45% C) in 0–50 cm depth; about 55% of this root production occured in the top 10 cm. Potential sources of error of the calculation method are discussed on the basis of the compartment flow model. Above-ground biomass production was 11.1 Mg ha^–1 in sole cropping and 13.6 Mg ha^–1 in alley cropping, of which 4.3 Mg ha^–1 were hedgerow prunings. The input of hedgerow root biomass into the soil was limited by the low root mass ofGliricidia as compared to other tree species, and by the decrease of live root mass of hedgerows and associated perennial weeds during the cropping season, presumably as a result of frequent shoot pruning.     

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 capacity in the component species of contour hedgerows: how important?

The choice of an appropriate hedgerow species is one of the most critical decisions in exploiting the value of a contour hedgerow system. The implications of hedgerow species with nitrogen (N)-fixation capacity on hedgerow-crop competition and crop productivity have been widely debated. We examined the agronomic significance of N-fixation by comparing the performance of species representing three classes of hedgerow vegetation: A nitrogen-fixing tree legumeGliricidia sepium), a non-nitrogen fixing tree (Senna spectabilis syn.Cassia spectabilis), and a forage grass (Pennisetum purpureum). The 4-year study investigated the hedgerow biomass and nutrient yields, and their relative effects on the performance of two annual crops commonly grown in alley farming systems, with emphasis on hedgerow-crop interference. The work was done on an Ultic Haplorthox (pH 4.8, organic C 1.9%, total N 0.18%).Senna produced 46% more pruning biomass on an annual basis than didGliricidia; N supplied to the alley crops was similar toGliricidia in the first year of observation, but 20–30% higher in the succeeding years. Upland rice and maize grain yields and total dry matter were unaffected by tree species, but the nitrogen-fixing tree exerted less competitive effects on the annual crops growing in adjacent rows. Grass hedgerows reduced maize yields 86% by the second year, indicating an unsustainable drawdown of nutrients and water. We conclude that hedgerow systems composed of a nitrogen-fixing tree did not exert significant advantages compared to a non-fixing tree species, and that factors other than N-fixation were more important determinants for the choice of hedgerow species.     

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.     

The effects of between-row (alley widths) and within-row spacings ofGliricidia sepium on alley-cropped maize in Sierra Leone

The effects of spacings between hedgerows (alley widths) and the spacings of trees within hedgerows ofGliricidia sepium on growth and grain yield of maize were investigated at Senehun in southern Sierra Leone. Four between-row spacings (2, 4, 6 and 8 m) were combined with three within-row spacings (0.25, 0.50 and 1.00 m) in a split block design. Maize, at densities of 20,000, 40,000 and 53,333 plants ha^–1, was established in the alleys and also as pure crops. N, P and K fertilizers were applied to all plots before pruning of the trees began. When pruning started, only the pure maize plots received fertilizer; prunings from the hedgerows were returned to the appropriate alleys in the other plots.Plots with the highest maize populations consistently gave the best yields before pruning started, but lower populations gave improved yields after pruning. Yields of maize increased with increasing alley widths before the start of pruning, after which the narrower alleys of 2 and 4 m outyielded the wider ones by almost double, probably because of the large amount of nutrients applied in prunings. Lack of light limited grain yields before the start of pruning, when there was some shading by the hedgerows. Alleys of 2–4 m wide, planted no closer than 0.50 m within rows, resulted in more than twice the yields of maize than in the 8-m alleys planted at 0.25 m within rows, once the hedgerows were well established and were being managed.     

Nitrogen mineralization and maize yields following application of tree prunings to a sandy soil in Zimbabwe

Despite the promotion of prunings as sources of nitrogen for crops, lack of synchronization between N mineralization from prunings and plant uptake remains a major limitation to the impact of prunings on crop yields. A laboratory and a field experiment were therefore carried out to determine the mineralization patterns of selected prunings and assess the potential that exists to improve synchrony by mixing prunings of different quality. The laboratory incubation experiment was conducted for 84 days to determine the C and N release patterns of prunings of different quality and the manipulation of the C and N mineralization trends by mixing prunings of different quality. High quality prunings were considered to be those with high CO₂ evolution rates and mineralize N rapidly. The % C and % N released in 84 days were highest for the high quality prunings of Tithonia diversifolia (70% and 30% respectively) and least for the low quality prunings of Flemingia macrophylla (25% and –5% respectively). The medium quality prunings of Acacia angustissima and Calliandra calothyrsus had similar proportions of released C and N (about 40% C and 10% N). Different mixtures of T. diversifolia with other species showed contrasting influence on C release and N mineralization. Most mixtures released less C than that predicted, but in contrast most mixtures released N at a rate either matching or above the predicted. The % N released was strongly correlated with the polyphenol protein binding capacity (r² = 0.53) and also with the % C released (r² = 0.62). The field experiment was done for two cropping seasons to determine the effects of the prunings of these species and their mixtures on maize grain yield. The prunings were added at 5 t ha^–1 and incorporated into the top 15 cm by hand hoeing in the first season and their residual effects were monitored in the second season. Medium and low quality prunings produced significantly (p < 0.05) higher maize grain yields in the first season ranging between 2.4 t ha^–1 and 3.4 t ha^–1 compared with T. diversifolia which produced 1.7 t ha^–1. This suggested better synchrony in N release and uptake by maize with medium and low quality prunings compared with high quality prunings. The only mixture that indicated improved synchrony was the mixture of T. diversifolia and C. calothyrsus. This study showed that mixing prunings of different quality produce different patterns of N mineralization, some of which were unexpected and had a potential for improving N synchrony.This revised version was published online in November 2005 with corrections to the Cover Date.     

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.     

Tree-crop interactions in hedgerow intercropping with different tree species and tea in Sri Lanka: 1. Production and resource competition

Incorporation of tree hedges along contours has been proposed as a means of reducing soil erosion and increasing soil fertility of tea (Camellia sinensis (L.) O. Kuntz) plantations on sloping terrain in high-rainfall zones of Sri Lanka. Tea yields in these hedgerow intercrops are determined by the balance between the positive (i.e., increased soil fertility) and negative (i.e., resource competition) effects of hedgerows. Therefore, the objectives of the present study were to determine: (a) the net effect of six different hedgerow species (Calliandra calothyrsus Meissner, Senna [Cassia] spectabilis (DC.) H. Irwin and Barneby, Euphatorium innulifolium (R.M. King and H. Rob.) H.B.K., Flemingia congesta Aiton F., Gliricidia sepium (Jacq.) (Kunth.) Walp. and Tithonia diversifolia (Hemsl.) A. Gray) on long-term tea yields; (b) the effect of incorporating hedgerow prunings as mulch; and (c) the relationship between hedgerow characteristics (i.e., biomass production, canopy and root growth) and tea yield variation. Tea yields, measured over one complete pruning cycle from October 1998 to September 2001, showed reductions relative to a sole tea crop under all hedgerow species except Euphatorium. The yield reductions ranged from 22 to 40%. Tea yields under Euphatorium showed increases up to 23% relative to the sole crop control. Addition of hedgerow prunings as mulch increased tea yields in all hedgerow intercrops. The yield increases ranged from 11 to 20%, with the highest being under Euphatorium. Tea yields showed a negative relationship (R²=0.38) with the pruned biomass of hedgerows. Limitation of environmental resources (e.g., water and light) and hedgerow characters which intensified resource competition (i.e., greater canopy lateral spread and height and greater root length densities, especially in the top soil layer) were responsible for observed tea yield reductions in hedgerow intercrops.     

Decomposition and phosphorus release from organic residues as affected by residue quality and added inorganic phosphorus

The combined use of organic residue and inorganic fertiliser-phosphorus (P) is appropriate in meeting both the short and long-term P requirement of crops. To assess the influence of added inorganic fertiliser-P on the processes of decomposition and P release from the residue and the relationships with quality, prunings of Gliricidia sepium, Leucaena leucocephela, Senna siamea, Acacia mangium and Paraserienthus falcataria were incubated without and with added inorganic fertiliser-P for 56 days. Soil was added only as inoculum. Decomposition rate and amounts of acid extractable-P (P release) were in the same order: G. sepium >S. siamea > L. leucocephela >P. falcataria > A. mangium. Unlike the other residues, A. mangium released no P despite the loss of half its mass during the 8 weeks of incubation. The residue P content correlated with P release. However, decomposition rate did not correlate with residue P content but with the lignin, polyphenol and cellulose content, and ratios to P. These ratios were negatively correlated with P release suggesting that lignin and polyphenol contents influence P release more when the residue-P content is low. Results suggest that rate of decomposition influences the release of P. The critical residue P content for P release was estimated to be 0.12% < P < 0.19%. Added P had no effect on decomposition and P release from the residues.     

Biomass production and root distribution of Gmelina arborea under an agrisilviculture system in subhumid tropics of Central India

A study of an agrisilviculture system comprising Gmelina arborea and soybean (Glycine max) was conducted in the subhumid region of Central India. Above- and below-ground biomass production and distribution of coarse and fine roots were studied in 4-year-old G. arborea, planted at a spacing of 2 × 2 m, 2 × 3 m, 2 × 4 m and 2 × 5 m. The total biomass varied from 10.89 Mg ha^–1 to 3.65 Mg ha^–1 depending on the tree density. Among the different tree components, stemwood contributed maximum biomass (54.3–79.4%), followed by branches and leaves. Root distribution pattern showed that most of the coarse roots were distributed in the top 40 cm of soil, whereas fine roots were concentrated in the top 20 cm. Coarse root biomass decreased with an increase in spacing. The spread of roots was asymmetrical in trees planted at 2 × 2 m and 2 × 3 m spacings, while it was symmetrical in trees planted at wide spacings. No significant difference was observed in the fine root biomass in different stands. The root:shoot ratio increased with an increase in spacing. Crop (soybean) growth and productivity varied significantly and it increased with a decrease in tree density. Soybean yield varied between 1.5 Mg ha^–1 to 2.1 Mg ha^–1. The role of root architecture of G. arborea trees on productivity of crops under agri-silviculture system is discussed.     

Implications of heterogeneity on procedures for estimating plant 15N recovery in hedgerow intercrop systems

Nitrogen flows in agroforestry systems can be quantified by applying excess ¹⁵N to one pool or part of the system and subsequentlymeasuring the quantity of ¹⁵N in other pools. Accurate quantifications depend on accurate determination of the mass, percentage N, and percentage ¹⁵N enrichment of each pool and past studies have mainly used physically isolated subplots to reduce variability. We thus assessed the within-plot and within-plant variability of ¹⁵N recovery by maize and by two hedgerow tree species, Gliricidia sepium and Peltophorum dasyrrachis, following applications of ¹⁵N-enriched materials to unbounded plots. We also assessed the potential for estimating total plant ¹⁵N recovery from a reduced set of samples, and for estimating N content allometrically from simple measurements. ¹⁵N uptake beyond 1 m radius was negligible in maize but recommended minimum inter-plot spacing for Gliricidia sepium and Peltophorum dasyrrachis was 8 m. Within-plant variation was also greater in trees than in maize. Calculating recovery from all crop plant components is not always necessary; the recovery of ¹⁵N in maize grain predicted 98.9% of variation in total plant ¹⁵N recovery. 89–95% accurate estimates of biomass and N content of maize plants and of regrowing tree shoots can be obtained non-destructively from height or basal shoot diameter respectively. ¹⁵N tracing techniques are potentially very useful for studies of competition and complementarity with respect to nitrogen uptake in agroforestry systems but they require unbounded plots and hence particular care in design and sampling procedures. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.