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.     

Nutrient requirements of exotic tree species in Zimbabwe

Pot and field experiments were conducted in the greenhouse and at three field sites (Marondera, Domboshawa and Makoholi) in Zimbabwe to examine the effects of soils and fertilizers on nutrient uptake and growth of 6 exotic tree species (Eucalyptus camaldulensis, E. grandis, E. tereticornis, Leucaena leucocephala, Casuarina cunninghamiana, and Acacia holosericea). Plant growth, N and P contents of all species were increased by the application of N, P, K and micronutrient fertilizers. The effect of individual nutrients (N, P, K and micronutrients) and their combination on Eucalyptus species was further investigated in a pot experiment using soil from Domboshawa. Eucalyptus species responded only to N application and no significant interactions were found between N and the other elements. Nutrient uptake results showed that E. camaldulensis and E. tereticornis removed more cations than the N-fixing trees but only in the fertilized treatments. L. leucocephala and C. cunninghamiana were higher in P, but no clear trends were observed for N. Plant growth and nutrient uptake by E. camaldulensis. C. cunninghamiana and A. holosericea were assessed in the field at the three sites. Plant species grown in the Marondera site had greater height and diameter at breast height (DBH) than those in the two other sites. These results followed trends in soil nutrient contents. The analysis of foliage revealed differences in the nutrient concentration of leaves from different trees, but no effect of site was found. The interrelationships between plant characteristics, soil and foliage nutrients were examined. In a pot experiment, mineral N was the only variable correlated with growth response and nutrient uptake, while in the field the only significant correlation was obtained with soil pH.     

Impact of residue quality on the C and N mineralization of leaf and root residues of three agroforestry species

A laboratory incubation experiment with ¹⁵N labeled root and leaf residues of 3 agroforestry species (Leucaena leucocephala, Dactyladenia barteri and Flemingia macrophylla) was conducted under controlled conditions (25 C) for 56 days to quantify residue C and N mineralization and its relationship with residue quality.No uniform relation was found between the chemical composition of the above and below residues. The leucaena and dactyladenia roots contained more lignin (8 and 26% respectively) and less N (2.0 and 1.0% respectively) than the respective leaves (2 and 13% lignin and 2.9 and 1.4% N, respectively), whereas the differences between the lignin and N contents of the flemingia leaves and roots were not significant (4.6 and 3.0% lignin and 2.63 and 2.68% N, respectively). The leucaena leaves contained more polyphenols than the roots (6.4 and 3.6%), while the polyphenol content of the leaves and roots of the other residues was similar (5.0 and 5.1% for dactyladenia and 4.0 and 3.5% for flemingia).Three patterns of N mineralization could be distinguished. A first pattern, followed by residues producing the highest amounts of CO₂, showed an initial immobilization of soil derived N, followed by a net release of both soil and residue derived N after 7 days of incubation. A second pattern, followed by the flemingia leaf residues which produced intermediate amounts of CO₂ and had an intermediate quality, showed no significant immobilization of soil derived N, and significant mineralization of residue N. A third pattern, followed by both low quality dactyladenia residues, showed a low release of residue derived N and a continued inmobilization of soil derived N.Residue C mineralization was significantly (p<0.05) correlated with the residue lignin content, C-to-N ratio, and polyphenol-to-N ratio. The proportion of residue N mineralized (immobilized) after 56 days of incubation was significantly correlated with the residue N content (p<0.01) and the C-to-N ratio (p<0.05). The relations were quadratic, rather than linear. The ratio of the proportion of residue N mineralized (immobilized) over the proportion of residue C mineralized after 56 days was highly significantly correlated with the lignin content (p<0.01) and C-to-N (p<0.001), lignin-to-N (p<0.01), polyphenol-to-N (p<0.01) and (lignin+polyphenol)-to-N ratios (p<0.01) in a linear way. This indicates that due to the low availability of the residue C, relatively less N is immobilized for the very low quality residues ((lignin+polyphenol)-to-N ratio: 29.7) than for the residues with a relatively higher quality ((lignin+polyphenol)-to-N ratios between 3.3 and 12.5).     

Assessment of genetic variability for N2 fixation between and within provenances of Leucaena leucocephala and Acacia albida estimated by 15N labelling techniques

Nitrogen fixed in 13 provenances of Acacia albida and 11 isolines of Leucaena leucocephala inoculated with effective Rhizobium strains was measured by ¹⁵N techniques and the total N difference method. In the test soil, on the average, L. leucocephala derived about 65% of its total N from atmospheric N₂ fixation compared to about 20% by A. albida. Significant differences in the percentage of N derived from atmospheric N₂ (% Ndfa) occurred, between provenances or isolines within species. The % Ndfa ranged from 37 to 74% within L. leucocephala and from 6 to 37 within A. albida; (equivalent to 20–50 mg N plant^–1 and 4–37 mg N plant^–1 for the two species over three months, respectively) and was correlated with the nodule mass (r=0.91). The time course of N₂ fixation of three selected provenances (low, intermediate and good fixers) was followed at 12 weekly intervals over a 36 week period. The % Ndfa of all provenances and isolines increased with time; and except for one of the L. leucocephala provenances, % Ndfa was similar within species at the 36 weeks harvest. There was a significant correlation between % Ndfa and the amount of N₂ fixed (r=0.96). Significant interactions occurred between provenances and N treatments and often growth of uninoculated but N fertilized plants was less variable than for inoculated unfertilized plants.     

The contribution of nitrogen by promiscuous soybeans to maize based cropping the moist savanna of Nigeria

Agronomic results indicate that maize grain yields generally are higher when the crop is planted following soybean than in continuous maize cultivation in the moist savanna agroecological zones of West Africa. Many factors have been hypothesized to explain this phenomenon, including enhanced N availability and the so-called `rotational effect'. There is, however, hardly any quantitative information on the residual N benefits of promiscuous soybeans to subsequent cereal crops grown in rotation with soybean. Three IITA promiscuous soybean breeding lines and two Brazilian soybean lines were grown in 1994 and 1995 at Mokwa in the southern Guinea savanna, Nigeria, to quantify the nitrogen contribution by soybeans to a succeeding crop of maize grown in rotation with soybean for two consecutive years, 1996 and 1997 using two methods of introducing ¹⁵N into soil (fresh ¹⁵N labelling and its residual ¹⁵N) and three maize cultivars (including one cultivar with high N use efficiency) used as reference plants. The nodulating soybeans fixed between 44 and 103 kg N ha^–1 of their total N and had an estimated net N balance input from fixation following grain harvest ranging from –8 to 43 kg N ha^–1. Results in 1996 and in 1997 showed that maize growing after soybean had significantly higher grain yield (1.2 – 2.3-fold increase compared to maize control) except for maize cultivar Oba super 2 (8644-27) (a N-efficient hybrid). The ¹⁵N isotope dilution method was able to estimate N contribution by promiscuous soybeans to maize only in the first succeeding maize crop grown in 1996 but not in the second maize crop in 1997. The first crop of maize grown after soybean accumulated an average between 10 and 22 kg N ha^–1 from soybean residue, representing 17–33% of the soybean total N ha^–1. The percentage ¹⁵N derived from residue recovery in maize grown after maize was influenced by the maize cultivars. Maize crop grown after the N-efficient hybrid cultivar Oba Super 2 (844-27) had similar ¹⁵N values similar to maize grown after soybeans, confirming the ability of this cultivar to use N efficiently in low N soil due to an efficient N translocation ability. The maize crop in 1997 grown after maize had lower ¹⁵N enrichment than that grown in soybean plots, suggesting that soybean residues contributed a little to soil available N and to crop N uptake by the second maize crop. The differential mineralization and immobilization turnover of maize and soybean residues in these soils may be important and N contribution estimates in longer term rotation involving legumes and cereals may be difficult to quantify using the ¹⁵N labelling approaches. Therefore alternative methods are required to measure N release from organic residues in these cropping systems.     

Persistence and recovery of introduced Rhizobium ten years after inoculation on Leucaena leucocephala grown on an Alfisol in southwestern Nigeria

Establishment of Leucaena leucocephala was poor at Ibadan (Transition forest-savanna zone) and Fashola (savanna zone, 70 km north of Ibadan) in southwestern Nigeria as a result of low soil fertility and the presence of only a few native rhizobia capable of nodulating it. Inoculation with L. leucocephala at these two locations in 1982 resulted in striking responses with Rhizobium strains IRc 1045 and IRc 1050 isolated from L. leucocephala grown in Nigeria. The persistence of inoculated effective Rhizobium strains after inoculation is desirable since it removes the need for reinoculation. Because of the perennial nature of L. leucocephala and its use in long-term alley farming experiments, we examined the persistence of inoculated rhizobial strains after inoculation, and their ability to sustain N₂-fixation and biomass production at Ibadan. In 1992, ten years after Rhizobium introduction, uninoculated, L. leucocephala fixed about 150 kg N ha^-1 yr^-1 or about 41% of total plant N compared to 180 kg N ha^-1 yr^-1 or 43% measured in 1982. Serological typing of the nodules using the Enzyme-Linked-Immunosorbent Assay (ELISA) and intrinsic resistance to the streptomycin test revealed that most of the nodules (96%) formed on L. leucocephala in 1992 were by Rhizobium strains IRc 1045 and IRc 1050, which were inoculated in 1982. Nodules were absent on uninoculated L. leucocephala grown on the adjacent field with no history of L. leucocephala cultivation. We conclude that the N₂ fixed by Rhizobium strains IRc 1045 and IRc 1050 persisted for many years in the absence of L. leucocephala and sustained effectively fixed N₂ which growth and yield of L. leucocephala after several years, thus encouraging a possible low-input alley farming system by smallholder farmers in Nigeria.     

Phosphorus benefits from grain-legume crops to subsequent maize grown on acid soils of southern Cameroon

We conducted field experiments over 2 years on two acid soils of southern Cameroon to test whether efficient uptake and use of phosphorus (P) from less available sources by grain legume genotypes could benefit subsequent rotational maize. We grew two crops each year. For the first crop we grew 4 genotypes of soybean and of cowpea, plus maize. For the second crop we grew maize. The first crops were fertilized with 0, 90 kg P ha⁻¹ as phosphate rock (PR) or 30 kg P ha⁻¹ as triple super phosphate (TSP). P application highly significantly increased shoot dry matter, P uptake, N₂ fixation and grain yields of the grain legumes with TSP generally more effective than PR. Two of the soybean and two of the cowpea genotypes were more efficient at using P. Only the P-efficient soybean and cowpea genotypes increased subsequent maize yields. Yields of the subsequent maize grown in rotation were significantly correlated with shoot P uptake for which the quantity of P applied with the crop residues of the pre-crop appeared to be a major factor. We also grew the grain legumes in nutrient solutions and measured organic acid-anion exudation from roots, root-surface phosphatase-activity, and root morphological characteristics. Enhanced exudation of organic acid anions from roots of P-deprived plants might have contributed to the P acquisition efficiency under field conditions of the P-efficient cowpea genotypes and one of the P-efficient soybean genotypes. A higher activity of root-surface acid phosphatase might have been important for the other P-efficient soybean genotype. The results show, that the potential positive rotational effect of cowpea and soybean on the acid, highly P-sorbing soils of southern Cameroon depends on breeding and using P-efficient genotypes when sparingly soluble and suboptimal rates of soluble P fertilizers are used. Section Editor: N. J. Barrow     

Senna siamea trees recycle Ca from a Ca-rich subsoil and increase the topsoil pH in agroforestry systems in the West African derived savanna zone

The functioning of trees as a safety-net for capturing nutrients leached beyond the reach of crop roots was evaluated by investigating changes in exchangeable cations (Ca, Mg, and K) and pH in a wide range of medium to long term alley cropping trials in the derived savanna of West Africa, compared to no-tree control plots. Topsoil Ca content, effective cation exchange capacity, and pH were substantially higher under Senna siamea than under Leucaena leucocephala, Gliricidia sepium, or the no-tree control plots in sites with a Bt horizon rich in exchangeable Ca. This was shown to be largely related to the recovery of Ca from the subsoil under Senna trees. The increase of the Ca content of the topsoil under Senna relative to the no-tree control treatment was related to the total amount of dry matter applied since trial establishment. The lack of increase in Ca accumulation under the other species was related to potential recovery of Ca from the topsoil itself and/or substantial Ca leaching. The accumulation of Ca in the topsoil under Senna had a marked effect on the topsoil pH, the latter increasing significantly compared with the Leucaena, Gliridia, and no-tree control treatments. In conclusion, the current work shows that the functioning of the often hypothesized ‘safety-net’ of trees in a cropping system depends on (i) the tree species and on (ii) the presence of a subsoil of suitable quality, i.e., clay enriched and with high Ca saturation.     

Evaluation of symbiotic properties and nitrogen contribution of mucuna to maize grown in the derived savanna of West Africa

The severity and increase of the Imperata cylindrica constraint as a weed, the decline of the traditional fallow systems as a means of soil fertility management and the lack of inorganic fertilizer appear to have created opportunities for adoption of mucuna (Mucuna pruriens) technology by smallholder farmers in some areas in the derived savanna of West Africa. What is not known, however, is the extent to which the establishment and N contribution of mucuna in these areas depend on symbiotic properties such as effective nodulation and mycorrhizal infection. Short term surveys carried out in 34 farmer's arable fields located in four different sites in the derived savanna, southern Benin, West Africa, together with results of greenhouse and field experiments showed that mycorrhizal infection rate of mucuma ranged from 2 to 31% and correlated positively with nodulation and shoot dry matter production. Nodulation occurred in 79% of the fields with numbers of nodules ranging from 0 to 135 plant^–1. Mucuna responded both to inoculation and N fertilizer in degraded soils but growth response depended on the rhizobia strains and mucuna varieties. Mucuna accumulated in 12 weeks about 313 kg N ha^–1 as either a sole crop or 166 kg N ha^–1 when mixed/intercropped with maize, respectively. Across all cropping systems it derived an average of 70% of its N from atmospheric N₂ (estimates made by the ¹⁵N isotope dilution method), representing 167 kg N ha^–1 per 12 weeks in the field. Mucuna interplanted with maize obtained a greater proportion of its nitrogen (74%) from fixation than did mucuna grown alone (66%) suggesting that competition for soil N influences the proportion of nitrogen fixed by mucuna. The total amount of N₂ fixed per hectare was, however, reduced significantly by intercropping mucuna with maize. A preceding mucuna crop provided a maize yield equivalent to 120 kg N kg ha^–1 of inorganic N fertilizer.     

Nodulation and growth response ofAllocasuarina andCasuarina species to phosphorus fertilization

To examine how soil phosphorus status affects nitrogen fixation by the Casuarinaceae —Frankia symbiosis,Casuarina equisetifolia and two species ofAllocasuarina (A. torulosa andA. littoralis) inoculated or fertilized with KNO₃ were grown in pots in an acid soil at 4 soil phosphate levels. InoculatedC. equisetifolia nodulated well by 12 weeks after planting and the numbers and weight of nodules increased markedly with phosphorus addition. Growth ofC. equisetifolia dependent on symbiotically fixed nitrogen was more sensitive to low levels of phosphorus (30 mg kg^–1 soil) than was growth of seedings supplied with combined nitrogen; at higher levels of phosphorus, the growth response curves were similar for both nitrogen fertilized and inoculated plants. The interaction between phosphorus and nitrogen treatments (inoculated and nitrogen fertilized) demonstrated that there was a greater requirement of phosphorus for symbiotic nitrogen fixation than for plant growth when soil phosphorus was low.WithAllocasuarina species, large plant to plant variation in nodulation occurred both within pots and between replicates. This result suggests genetic variation in nodulation withinAllocasuarina species. Nodulation ofAllocasuarina species did not start until 16 weeks after planting and no growth response due toFrankia inoculation was obtained at the time of harvest. Addition of nitrogen starter is suggested to boost plant growth before the establishment of the symbiosis. Growth ofAllocasuarina species fertilized with nitrogen responded to increasing levels of phosphorus up to 90 mg P/kg soil after which it declined by 69% forA. littoralis. The decrease in shoot weight ofA. littoralis, A. torulosa, C. equisetifolia andC. cunninghamiana at high phosphorus was confirmed in a sand culture experiment, and may be atributable to phosphorus toxicity.