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 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.     

Effect of successive cuttings on uptake and partitioning of 15N among plant parts of Leucaena leucocephala

Summary We studied the effect of three successive cuttings on N uptake and fixation and N distribution in Leucaena leucocephala. Two isolines, uninoculated or inoculated with three different Rhizobium strains, were grown for 36 weeks and cut every 12 weeks. The soil was labelled with 50 ppm KNO₃ enriched with 10 atom % ¹⁵N excess soon after the first cutting. Except for the atom % ¹⁵N excess in branches of K28 at the second cutting, both the L. leucocephala isolines showed similar patterns of total N, fixed N₂, and N from fertilizer distribution in different parts of the plant at each cutting. The Rhizobium strain did not influence the partitioning of ¹⁵N among the different plant parts. Significant differences in ¹⁵N enrichment occurred in different parts. Live nodules of both isolines showed the lowest atom % ¹⁵N excess values (0.087), followed by leaves (0.492), branches (0.552), stems (0.591), and roots (0.857). The roots contained about 60% of the total plant N and about 70% of the total N derived from fertilizer over the successive cuttings. The total N₂ fixed in the roots was about 60% of that fixed in the whole plant, while the shoots contained only 20% of the fixed N₂. We conclude that N reserves in roots and nodules constitute another N source that must be taken into account when estimating fixed N₂ or the N balance after pruning or cutting plants. ¹⁵N enrichment declined up to about fivefold in the reference and the N₂-fixing plants over 24 weeks following the ¹⁵N application. The proportion and the amounts of N derived from fertilizer decreased, while the amount derived from N₂ fixation increased with time although its proportion remained constant.     

Phosphorus requirements and nitrogen accumulation by N2-fixing and non-N2-fixing leguminous trees growing in low P soils

The variation in P uptake and use efficiency and N accumulation by Gliricidia sepium (N₂-fixing tree), Senna siamea and S. spectabilis (leguminous non-N₂-fixing trees) were examined in the field at Fashola (savanna zone), southwestern Nigeria, using four P rates, 0, 20, 40 and 80 kg P ha^-1. Growth of G. sepium and S. spectabilis responded to P application at 24 weeks after planting (WAP) and average yield increases of 58% and 145% were observed by the application of 40 kg P ha^-1 for the two species, respectively. Such a P response was not found in S. siamea at 24 WAP and for any of the species at 48 WAP. G. sepium accumulated more P (on average 162%) than S. siamea and S. spectabilis at 24 WAP and had greater root length and a higher percentage of mycorrhizal infection. However, at 48 WAP S. siamea had 2.5 times more P than G. sepium. Differences in the physiological P use efficiency (PPUE) between G. sepium and the non-N₂-fixing trees were significant at the 0 P level, being higher for S. siamea (average, 0.61 g shoot mg^-1 P) than for G. sepium (0.27 g shoot mg^-1 P). G. sepium had a consistently lower atom % ¹⁵N than S. spectabilis, while that of S. siamea for most of the time did not differ from that of G. sepium. The reference plant affected N₂ fixation extimates, with negative values and a higher variability (CV 60%) associated with S. siamea than with S. spectabilis (CV<20%). Consequently, S. spectabilis was selected as a better reference plant for measuring N₂ fixation in G. sepium. G. sepium fixed on average 35% and 54% of its N at 24 and 48 WAP, respectively. Except at the lowest P rate, percentage and amount of N fixed were not generally enhanced by P application.     

Time-course of dinitrogen fixation of promiscuous soybean cultivars measured by the isotope dilution method

 Soybean cultivars capable of nodulating with indigenous Bradyrhizobium spp. have been developed by the International Institute of Tropical Agriculture (IITA) and national programs in Africa in order to avoid artificial inoculation by resource-poor farmers in Africa. The current selection procedure for enhanced N₂ fixation is based on an assessment of nodule formation which does not directly quantify the proportions of crop N derived from the atmosphere. We have monitored N accumulation patterns and N₂ fixation in nine promiscuous soybean cultivars with different maturity periods, using the ¹⁵N dilution technique. Nodule development generally peaked at the early podfill stage for all cultivars except Tgx 1519-1D and Tgx 1447-2D in which it continued to increase. The proportion of crop N derived from fixation (%NDFA) ranged between 51% and 67%, 77% and 84%, and 66% and 73% at full bloom, early podfill, and physiological maturity stages, respectively. Total N accumulation increased in all soybean genotypes with increasing plant age. Significant correlations (P<0.001) were established between nodule weight and %NDFA, even though this did not explain the relationship between nodule development and N₂ fixation in cultivars such as Tgx 1519-1D. Promiscuous soybean cultivars retained between 10% and 19% of total N accumulated at the final harvest, in belowground biomass. Our results indicated that these soybean cultivars can derive substantial proportions of plant N from N₂ fixation in soils where compatible indigenous bradyrhizobia populations are adequate and effective. Also, we have substantiated the claims that qualitative nodulation parameters currently used to select varieties with a high N₂ fixation capacity need to be validated with other measurements of N₂ fixation. Received: 5 November 1998     

Leaf quality of selected hedgerow species at two canopy ages in the derived savanna zone of West Africa

Prunings in hedgerow intercropping systems are a potential source of organic matter and their quality is an important characteristic driving decomposition and nutrient release. To determine the potential range of residue characteristics and the impact of canopy age on those, selected characteristics of Leucaena leucocephala, Gliricidia sepium, and Senna siamea hedgerow leaf residues were determined for young (5–10 weeks) and old (39–47 weeks) hedgerow canopies in six sites, representative in terms of soil for the derived savanna zone of West-Africa. The N content of the Leucaena and Senna residues decreased with age (from 4.8 to 3.9% and from 4.1 to 3.0%, respectively, while the N content of the Gliricidia residues remained constant (3.9%). The P content of all species decreased with age (from 0.30 to 0.20%, on average). The lignin content of the Leucaena residues increased, and their polyphenol content decreased with age, while for the Senna residues only an increase in polyphenol content with canopy age was evident. Neither the lignin nor the polyphenol content of the Gliricidia leaf residues was markedly affected by canopy age. For all species, the P and ADF content were positively correlated with the ash and lignin content, respectively. The N content of the Leucaena and Senna residues was positively correlated with their P content while the lignin content of the same species was negatively correlated with the polyphenol content. Because of the low variation in certain residue characteristics between sites, the presented data and equations could potentially be used to reasonably estimate those characteristics if no other information is available. For other characteristics, such as polyphenol contents, measurements are preferable in view of the relatively high variation encountered.This revised version was published online in November 2005 with corrections to the Cover Date.     

Nodulation and estimation of symbiotic nitrogen fixation by herbaceous and shrub legumes in Guinea savanna in Nigeria

Twelve herbaceous and shrub legume species were grown in pot and field experiments in five sites representing three agroecological zones in moist savanna in Nigeria. The objectives were to: (1) assess natural nodulation of the legumes and characterize their indigenous rhizobia, (2) determine their need for rhizobia inoculation and (3) estimate the amount of N₂ fixed by each of these legumes. At 4 weeks after planting (WAP), Crotolaria verrucosa was not nodulated at any of the sites while Centrosema pascuorum had the highest number of nodules in all sites. At 8 WAP, all legumes were nodulated, with Mucuna pruriens having the least number of nodules and Stylosanthes hamata the highest. The number of nodules, however, was inversely correlated to the mass of nodules. Significant differences in nodulation of the legume species grown in the field also occurred between and within sites. Mucuna pruriens and Lablab purpureus produced more shoot and nodule biomass than the other legumes in all sites. Growth of most of these legumes responded to fertilizer application, except for C. verrucosa and Aeschynomene histrix. Except for C. verrucosa, average proportion of N₂ fixed was about 80% and this was reduced by about 20% with N fertilizer application. The majority of rhizobia isolates (60%) were slow growing, belonging to the Bradyrhizobia spp. group. Selected rhizobia isolates evaluated on Cajanus cajan, C. pascuorum, M. pruriens and Psophocarpus palustris varied from ineffective to highly effective in Leonard jar conditions. However, only growth of M. pruriens responded to inoculation in potted soils, whereas it was lower than that obtained with N fertilizer application. This indicated the need to screen more rhizobia in order to improve N₂ fixation and growth of legume species such as M. pruriens when it is introduced in soils deficient in N.     

Field measurements of nitrogen fixation in leguminous trees used in agroforestry systems: Influence of 15N-labeling approaches and reference trees

Appropriate ¹⁵N-labeling methods are crucial for estimating N₂-fixation in trees used in agroforestry systems. A 4-year field experiment was conducted on an Alfisol in Southwestern Nigeria to compare the estimates of N₂ fixed in Leucaena leucocephala, using two non-N₂-fixing leguminous trees, Senna siamea and S. spectabilis, as reference plants and three different methods of introducing ¹⁵N into soil. The atom % ¹⁵N uptake pattern (as reflected in the leaves) was identical in both N₂- and non-N₂-fixing tree species irrespective of the ¹⁵N-application method. There was a significant decline in atom % ¹⁵N excess in the leaves of L. leucocephala (from 0.266 to 0.039), S. siamea (0.625 to 0.121), and S. spectabilis (from 0.683 to 0.118) from the first sampling 12 months after planting and the second sampling 18 months after sampling. From the second harvest in 1991 until the end of the experiment (fifth) harvest in 1993, however, the atom ¹⁵N % excess decline in leaves of the three species was less pronounced and depended on the method of ¹⁵N application. In those plants to which the tracer was applied once at planting, the ¹⁵N decline was steady between the second and the last prunings. In the split-application treatment, the atom ¹⁵N % excess increased slightly at the third pruning and decreased during the subsequent two prunings. The reference tree and the method of ¹⁵N application influenced the estimated proportion of N derived from atmospheric N₂ by L. leucocephala, calculated as 73 and 64%, corresponding to 119 and 98 kg N ha^-1 of N₂ fixed per 6 months, when S. spectabilis and S. siamea were used as reference trees, respectively. The approach by which ¹⁵N-labeled fertilizer was applied to the soil in three splits gave slightly higher estimates of N derived from the atmosphere but this was of little agronomic significance because total N₂ fixed was similar for all methods.     

Effects of Mulch from 14 Multipurpose Tree Species (MPTs) on Early Growth and Nodulation of Cowpea (Vigna unguiculata L.)

Leaf extracts and mulch from 14 MPTs were used to study their effects on germination, initial growth and nodulation of cowpea. Aqueous leaf extracts of , , and full concentrations were added to cowpea seeds placed in Petri dishes. In the pot experiment, cowpea was planted in plastic pots filled with 1200 g of soil, and mulch was either surface-applied or incorporated into the substrate at rates equivalent to 5, 10 and 20 tons · ha⁻¹. Leaf extracts from Gliricidia sepium, Senna siamea, Milletia thoningii, Grewia pubescens and Tetrapleura tetraptera reduced cowpea germination significantly at concentrations above 25 % of the full concentration 7 days of incubation. Root length was also strongly reduced by extracts from Gmelina arborea, Senna siamea, Alchornea coordifolia, and Tetrapleura tetraptera. These inhibiting effects were also observed in the pots. Mulch incorporation significantly reduced growth and nodulation of cowpea in the pots. These reductions were more pronounced with the incorporation of mulch from fast decomposing trees such as Leucaena leococephala, Gliricidia sepium and Grewia pubescens. Although Leucaena leucocephala and Gliricidia sepium released much nitrogen, they strongly reduced cowpea growth when incorporated. This may suggest that other reasons than N immobilization caused growth reduction when the plant residues were applied to cowpea.     

Assessment of soil factors limiting growth and establishment of Mucuna in farmers' fields in the derived savanna of the Benin Republic

Mucuna pruriens was been used to control Imperata cylindrica and improve soil fertility in maize and cassava cropping systems in the derived savanna of the Benin Republic, West Africa. However, field observations showed that Mucuna had poor establishment in some farmers' fields. This could be due in part to the poor symbiotic effectiveness of Mucuna and/or its poor nutrition because of mineral deficiencies in the soil. A short-term survey was carried out in 34 farmers' fields located in four different sites (Zouzouvou, Eglime, Tchi, and Niaouli) in the derived savanna to assess the natural nodulation and mycorrhizal infection of Mucuna. This survey was followed by a nutrient-omission trial conducted in a pot experiment using soil collected from two groups of farmers' fields at Zouzouvou where Mucuna had poor establishment. Mycorrhizal infection ranged from 2 to 31% and correlated positively with nodulation and shoot dry matter production of plants grown only in one site at Zouzouvou. The number of rhizobia ranged between <0.05 (near the detection limit) and 15 cells g^-1 soil depending on the plot history and the fields. Nodulation occurred in 79% of the fields with numbers of nodules ranging from 0 to 135 plant^-1. The nutrient-omission trial showed that when N and P were absent in the complete fertilizer treatment, biomass production decreased significantly, on average by 69% (N) and 33% (P). Mg, S, K and micronutrient deficiencies did not reduce significantly the biomass production in the two groups. However, N fertilizer applied additionally each week to some treatments drastically reduced Mucuna nodulation. Strategies to enhance Mucuna establishment and growth are discussed.