Nitrogen dynamics in maize-based agroforestry systems as affected by landscape position in southern Malawi

In Malawi, agroforestry is very promising for N replenishment; however, there are still large variations in the performance of these agroforestry technologies on farmers’ fields. A study was conducted on-farm to determine the influence of three landscape positions on N dynamics in maize (Zea mays L.)-based agroforestry systems. The agroforestry systems were relay fallow using Sesbania sesban (L.) Merr or Tephrosia vogelii (Hook F.), simultaneous fallow using Gliricidia sepium(Jacq.) Walp., and maize without trees as a control. Sesbania was superior to other systems in the bottom slope, producing the highest tree biomass (1,861 kg ha⁻¹), whereas, gliricidia gave the highest tree biomass production in the mid-slope (2,147 kg ha⁻¹) and upper slope (1,690 kg ha⁻¹). Preseason inorganic N, maize flag leaf N concentration, maize total N uptake and maize yields followed a similar trend to tree biomass production with tree-based cropping systems exhibiting higher productivity (P < 0.05) than the cropping systems without trees. Nitrogen leaching from gliricidia agroforestry systems was lower than in the other agroforestry systems across all landscape positions as evidenced by 17% lower amounts (P < 0.05) of inorganic N adsorbed to ionic exchange resin membranes at 60 cm soil depth most likely due to the permanent root system of gliricidia. The difference between δ¹⁵N values of the trees and the soil did not change between landscape positions suggesting that if the leguminous trees fixed atmospheric N₂, the proportion of total N uptake was identical at all locations. We concluded that landscape positions have a significant effect on tree performance with sesbania remarkably adapted to the bottom slope, gliricidia to the mid-slopes and tephrosia fairing similar in both the bottom slope and mid-slopes.     

Microcosmic study of soil microarthropod and earthworm interaction in litter decomposition and nutrient turnover

A microcosm experiment was set up under laboratory conditions and verified under field conditions with the objective of investigating the interaction of soil microarthropods and earthworms in litter decomposition, nutrient release, and uptake by maize crop. The treatments included: soil alone (control), soil with leaf litter (Senna siamea leaves), soil with leaf litter and soil microarthropods, soil with leaf litter and earthworms (Hyperiodrilus africanus), and soil with litter and both of the soil faunal groups. After an 8-week incubation period, the amounts of litter decomposed and N, P, K, Ca, and Mg released followed the order: with microarthropods and earthworms > with earthworms > with microarthropods > no faunal addition. The presence of microarthropods and earthworms also increased the net level of mineral N in the incubated soil. The additive roles of soil microarthropods and earthworms were observed on decomposition and nutrient release. Such faunal interactions resulted in an increased N uptake by maize in the incubated soil. Despite their lower biomass, soil microarthropods contributed significantly to nutrient turnover in the presence of earthworms. This study emphasizes the need to quantify and devise ways of controlling and regulating the abundance and activities of soil fauna for effective nutrient cycling and, consequently, for better crop yields in low-input tropical agricultural ecosystems.     

Litter decomposition, microbial biomass and activity of soil organisms in three agroforestry sites in central Amazonia

Soil organisms play a central role in the decomposition of organic matter. The activity of soil organisms was comparatively examined in three experimental sites in central Amazonia (Brazil): a peach palm monoculture (Bactris gasipaes) a, rubber tree plantation (Hevea sp.), and an agroforestry system (four tree species planted in rows, the space between covered by upcoming secondary vegetation). The overall decomposition rates in the systems and the role of different groups of soil organisms (macrofauna, mesofauna, microflora) were studied with leaf litter (Vismia guianensis) enclosed in litter bags. Microbial respiration and biomass (SIR method) in litter and soil were measured (IRGA). Microbial respiration in all sites decreased in the gradient litter > topsoil (0–5 cm) > soil at 5–15 cm. The highest decomposition rate was always observed in the litter bags of coarse mesh size, pointing to the crucial role of the macrofauna in maintaining a high decomposition rate of the organic material in all systems. The Hevea (k = 3.4) and the Bactris plantation (k=3.1) both showed the highest decomposition rates, followed by the polyculture system (k=1.9). The Bactris plantation also had the highest level of microbial respiration and biomass in litter and soil. We discuss these findings in the light of data on rainfall, pH and canopy closure. They suggest that microclimate is a more important factor than biomass in determining litter decomposition rates and activity of soil organisms at these sites.     

Arbuscular Mycorrhizal Fungi and Plant Chemical Defence: Effects of Colonisation on Aboveground and Belowground Metabolomes

Arbuscular mycorrhizal fungal (AMF) colonisation of plant roots is one of the most ancient and widespread interactions in ecology, yet the systemic consequences for plant secondary chemistry remain unclear. We performed the first metabolomic investigation into the impact of AMF colonisation by Rhizophagus irregularis on the chemical defences, spanning above- and below-ground tissues, in its host-plant ragwort (Senecio jacobaea). We used a non-targeted metabolomics approach to profile, and where possible identify, compounds induced by AMF colonisation in both roots and shoots. Metabolomics analyses revealed that 33 compounds were significantly increased in the root tissue of AMF colonised plants, including seven blumenols, plant-derived compounds known to be associated with AMF colonisation. One of these was a novel structure conjugated with a malonyl-sugar and uronic acid moiety, hitherto an unreported combination. Such structural modifications of blumenols could be significant for their previously reported functional roles associated with the establishment and maintenance of AM colonisation. Pyrrolizidine alkaloids (PAs), key anti-herbivore defence compounds in ragwort, dominated the metabolomic profiles of root and shoot extracts. Analyses of the metabolomic profiles revealed an increase in four PAs in roots (but not shoots) of AMF colonised plants, with the potential to protect colonised plants from below-ground organisms.     

Decomposition and nutrient release from pruning residues of two indigenous agroforestry species during the wet and dry seasons

The decomposition of leaves from Cordia africana Lam. and Albizia gummifera G. F.Gmel was investigated during the wet and dry seasons at Wondo Genet (Ethiopia). Litterbags of leaves were buried in soils under farmland and shaded-coffee agroforestry systems. Residual matter was recovered after 4, 8, 12, and 16 weeks and analysed for nitrogen (N), phosphorus (P), potassium (K), cellulose, lignin, soluble polyphenol and condensed tannin content. Mass-loss and release of N, polyphenols and condensed tannins were greater from Albizia leaves than from Cordia leaves, suggesting that a high polyphenol content does not necessarily retard decomposition. The rates of mass loss and release of the majority of leaf constituents were considerably faster during the wet season than during the dry season. Lignin decomposition, however, proceeded more rapidly during the dry season, and no significant seasonal differences were observed for polyphenol decomposition. The decomposition kinetics of most leaf components during the wet season were best described by a single-exponential model, but a quadratic model provided the best fit during the dry season. Initial leaf chemistry and season were important decomposition factors, while land-use effects were negligible. However, land-use effects showed distinct seasonal differences, with leaf litter decomposing more rapidly in soil under shaded-coffee than under farmland management, especially during the wet season. This study also demonstrated that polyphenol content does not show the predictive effects it has been attributed to have and that other constituents, such as condensed tannins, would be better suited for this purpose.     

Nutrient release during decomposition of leaf litter in a peach (<Emphasis Type="Italic">Prunus persica L.</Emphasis>) orchard

The decomposition of plant residues has a pivotal role in carbon and nutrient cycles in agricultural ecosystems, where it can contribute to the sustainable management of the crops. In this paper we report a study on the release of C and nutrients during the decomposition of peach abscised leaves in northern Italy. Litter bags containing a representative amount of leaf material were installed under tree canopies either on the soil surface under open field conditions or on the soil surface in pots under partially controlled conditions. Potted leaf litter was pre-labelled with ¹⁵N. We observed that in 3 years, peach leaves lost 85% of their initial mass mostly attributable to cellulose decomposition, while new lignin or lignin-like compounds were synthesised during the first stages of the decomposition process. Nutrient dynamics differed depending on the considered element. Nitrogen and sulphur were initially immobilized into the litter to be released only starting after 44 weeks from the beginning of the decomposition. Potassium and magnesium were rapidly released in the winter following leaf abscission, reaching an amount that remained constant up to the end of the trial. Calcium and phosphorus release was slow, but constant throughout the three-year study period. With the only exception of Mg, 80–90% of initial amounts of mineral elements had released from decomposing peach leaves after 3 year from leaf abscission. Since in mature stands leaf litters of different ages coexist on the soil surface 80–90% of the nutrients contained in the abscised leaves are expected to return annually to the soil and potentially be available for subsequent root and/or microbial uptake.     

Litter fall, litter stocks and decomposition rates in rainforest and agroforestry sites in central Amazonia

The sustainability of agroforestry systems in Amazonia was assessed from their litter dynamics and decomposition. Litter fall and litter stocks were determined from July 1997 to March 1999 in four sites in central Amazonia: a primary rainforest, a 13-year-old secondary forest, and two sites of a polyculture forestry system which consisted of four planted tree species of commercial use amidst upcoming secondary growth. The average annual litter fall in the undisturbed primary rainforest (FLO) was 8.4 t ha^–1 year^–1, which is within the range of litter fall in other rainforests in the region. It was similar in one of the two polyculture sites (8.3 t ha^–1 year^–1), but lower in the secondary forest and in the second polyculture site. In the litter fall in secondary forest and agroforestry sites, the leaf portion was higher (76–82% of total litter fall) than in FLO, due to reduced fine matter and wood fall. Leaf litter fall variability was much lower in the plantation sites than in the forests, which is explained by the much more homogeneous stand structure of the plantations. The quality of the produced litter, measured as C/N ratio, differed significantly between the primary forest site and one polyculture and the secondary forest site. The cumulative input of nitrogen through litter fall was 144 kg ha^–1 year^–1 in FLO, and 91–112 kg ha^–1 year^–1 in the polycultures and the secondary forest. Litter fall was not correlated with soil parameters, but had a significant linear regression with canopy closure. For the primary rainforest, litter fall was also (inversely) correlated with monthly rainfall. Litter fall was higher in the first year (1997–1998; an El Niño period) than in 1998–1999. Litter stocks on the forest floor were highest in the secondary forest (24.7 t ha^–1), and much lower in the polyculture sites (15.1–16.2 t ha^–1) and the primary forest (12.0 t ha^–1). There were no differences in the relative N content (C/N ratio) of the litter stocks between the sites, but the larger stocks led to higher absolute N contents in the litter layer in the secondary forest. From the monthly values of litter stocks (S) and litter fall (P), the decomposition coefficient k _e=P/S was calculated, which was, on average, highest for the primary forest (0.059), followed by the polyculture systems (0.040–0.042), and by the secondary forest (0.024). Thus, due to low decomposition rates, the secondary forest site showed large litter accumulations in spite of a relatively low litter fall. In contrast, the primary forest showed high litter fall but low stocks, due to high decomposition rates. The decomposition coefficients of the polyculture systems ranged between the primary and the secondary forest. The reduced decomposition rates in the man-managed agroecosystems indicate quantitative and/or qualitative changes in the decomposer communities of these systems that lead to a higher build-up of litter stocks on the forest floor. However, the decomposer systems in the polyculture sites still were more functional than in the site of non-managed secondary growth. Thus, from a soil biological viewpoint, ecologically sustainable low-input agroforestry in Amazonia will benefit from the application of these polyculture systems.     

Responses of plant traits of four grasses from contrasting habitats to defoliation and N supply

The objective of the study was to identify specific plant traits determining adaptation of grass species to defoliation and N availability, and thus having a major impact on species dynamics, primary productivity, and on nutrient cycling in grassland ecosystems. It was specifically examined whether the response of species to defoliation is related to their plasticity in leaf growth and in leaf growth zone components, and whether the response of species to nitrogen is related to their plasticity in root morphology and subsequent N acquisition, and to N losses through leaf senescence. The study was conducted on L. perenne and D. glomerata, two grazing tolerant species from fertile habitats, and on F. arundinacea and F. rubra, two less grazing tolerant species from less fertile habitats. Plants were subjected to repeated defoliation at three cutting heights under both high N and low N supply. Biomass allocation, leaf elongation, characteristics of the leaf growth zone (height and relative growth rate), and root morphology, N uptake and N losses through leaf senescence were evaluated. Under high N supply, L. perenne and D. glomerata showed the greatest tolerance to defoliation, due to a large plasticity in the height of the leaf growth zone and due to compensatory growth, either within the leaf growth zone or between growing leaves. Under low N supply, F. rubra was the only species with the ability to develop a more branched root system and a greater length of tertiary roots than under high N. As a consequence, under low N supply F. rubra had a higher specific N uptake and a higher growth rate than the other species. This slow growing species also showed a higher nitrogen allocation to dead leaves and subsequently a higher potential N loss to leaf litter.     

How does plant leaf senescence of grassland species influence decomposition kinetics and litter compounds dynamics?

The influence of litter quality on plant litter decomposition rates is a crucial aspect of the soils C cycle. In grassland ecosystems, leaf litter, which is not removed either by herbivores or by mowing, returns to soil after the senescence process (brown litter). In grassland managed by mowing, another significant proportion of litter returns to the soil before senescence through harvesting losses (green litter). We hypothesized that changes in leaf tissue quality due to the senescence process would lead to contrasting decomposition dynamics of brown litter compared to green litter. Our conceptual approach included the monitoring of decomposition of green (fresh leaves) and brown litter (dead leaves, still attached to the plant) of three different grassland species (Lolium perenne, Festuca arundinacea and Dactylis glomerata) during a 1 year field incubation. After 0, 2, 4, 20 and 44 weeks, we retrieved the litterbags and analysed the remaining material for carbon and nitrogen content and stable isotope composition. Additionally, we determined the lignin content and composition by CuO oxidation and the non-cellulosic neutral carbohydrate content and composition after TFA hydrolysis. As expected, green litter, being higher in N and soluble compounds, while showing a lower C:N ratio and lower lignin contents compared to brown litter, was degraded at a higher rate. Carbon decomposition kinetics suggests that both leaf litter types consist of two pools with contrasting turnover times. The size of the active pool was related to the initial content of soluble plant litter compounds and the size of the recalcitrant pool was related to the lignin to N ratio of initial plant material. More lignin was lost from green litter compared to brown litter. P-coumaryl-type lignin units were decomposed at a higher rate than vanillyl and syringyl units. Total non cellulosic polysaccharide content showed little changes for both litter types. In contrast, the ratios of hexoses/pentoses (C6/C5) and desoxy sugars/pentoses (desoxy/C5) increased during decomposition of green litter only. This is an indication for an increasing contribution of microbial derived compounds being consistant with the higher decomposition rate of this material. Our results showed that grassland management (grazing versus mowing) could influence soil carbon sequestration through different proportions of green and brown litter returned to soil.     

Role of allelopathy as a possible factor associated with the rising dominance ofBunias orientalis L. (Brassicaceae) in some native plant assemblages

Leaf extracts ofBunias orientalis were shown to inhibit seed germination of a variety of cultivar plant species and of species cooccurring withB. orientalis in the field. Root exudate solutions and leaf litter leachates ofB. orientalis were tested for their allelopathic activity using seedling growth assays. Additionally, in comparative seedling growth assays soil cores removed from denseB. orientalis stands were tested bimonthly for elevated allelopathic effects. The impact of root exudates on seedling growth was generally weak and varied between species. Similar results were obtained for the effect ofB. orientalis leaf litter leachates on seedlings grown in sand culture relative to the effect of leaf litter leachates of a plant species mixture. When soil as a growth substrate was used, no consistent differences in seedling growth were obtained between the two litter leachate treatments. In the soil core experiment seedlings grown in soil cores collected from a denseB. orientalis stand unexpectedly showed better performance than seedlings grown in soil cores collected from a nearby mixed plant stand withoutB. orientalis, at least in early spring and late autumn. Predominating nutrient effects are, therefore, assumed to conceal a potentially increased allelopathic effect of soil beneath denseB. orientalis stands. It is concluded that other factors than allelopathy must be investigated to explain the rapid establishment of dense stands of this alien plant species.     

Estimation of N2 fixation in Desmodium ovalifolium from the relative ureide abundance of stem solutes: Comparison with the 15N-dilution and an in situ soil core technique

Many, but not all, legumes of tropical origin, transport fixed N from the nodules to the shoot tissue in the form of ureides, and the mineral N absorbed from the soil is principally transported in the form of nitrate. The analysis of stem xylem sap, or hot-water extracts of stem tissue, for ureide and nitrate has been used successfully to quantify BNF contributions to several grain legumes and more recently to some shrub and forage legumes. The objective of this study was to investigate the application of this technique to the quantification of the contribution of BNF to the forage legume Desmodium ovalifolium by comparing the relative ureide abundance (RUA) of stem extracts of this plant with simultaneous estimates of BNF obtained using the ¹⁵N isotope dilution technique. The first experiment was performed in pots of soil, taken from a grazing study, amended with ¹⁵N-labelled organic matter at four different application rates. The ureide concentration in the stem extracts reflected the changes in BNF activity during plant growth and the RUA was closely correlated with the proportion of N derived from BNF as determined from the ¹⁵N technique (r ² = 0.86 and 0.88 for inoculated and non-inoculated plants, respectively). The use of a calibration curve derived from a previous study where the same legume was fed increasing concentrations of ¹⁵N labelled nitrate in sand/vermiculite culture, resulted in an over-estimation of the BNF contribution which may have been due to a significant uptake of ammonium from this acidic soil. The second experiment was performed in field plots and a good agreement was found between the estimates of BNF derived from using the ureide and ¹⁵N dilution techniques at two harvests six months apart. The uptake of soil N by the D. ovalifoliumand two forage grasses (Brachiaria humidicola and Panicum maximum) was estimated using an in situ soil core technique, and, while the uptake of N by the grasses was successfully estimated, this technique underestimated the N derived from the soil by the legume as determined by the ureide and ¹⁵N dilution techniques.     

The Plant Growth Inhibitor Nagilactone Does Not Work Directly in a Stabilized Podocarpus nagi Forest

Nagilactones isolated from Podocarpus nagi are known for poisonous physiological activities to organisms. To clarify the allelopathic potential of nagilactone to wild plants, the seasonal changes in nagilactone contents in throughfall, leaf litter, and soil were monitored monthly in a podocarp forest at Mt. Mikasa, Nara City, central Japan. The average annual nagilactone concentrations in throughfall, leaf litter, and surface soil at 0–10 cm depth were 4.7 × 10^–12 g/ml, 3.8 × 10^–4 g/g and 3.1 × 10^–7 g/g, respectively. The nagilactone flux from canopy leaves to the forest floor via throughfall was 5.2 × 10^–2 g/ha/yr, which was far smaller than that via leaf litterfall: 765.3 g/ha/year. The disappearance of nagilactones from leaf litter and the nagilactone accumulation in the Ao layer and mineral soil were also studied. Nagilactone in leaf litter rapidly disappeared and its relative disappearance rate was 5.8/yr. Nagilactone concentration in mineral soil also decreased rapidly with an increase in soil depth and could not be detected in soil at 40–100 cm depth. The total mass of nagilactones was 177.3 g/ha in the Ao layer and 105.0 g/ha in mineral soil. These quantities in nagilactone dynamics give a very rapid turnover time of 0.37 year (4.4 months) in a soil nagilactone pool, implying the possibility of the reabsorption of nagilactone by P. nagi, while the allelopathic effects of nagilactone to other plants in the podocarp forest are questionable.     

Damage-induced alkaloids in tobacco: Pot-bound plants are not inducible

Field-grown wild tobacco plants (Nicotiana sylvestris) were subjected to a defoliation regime designed to mimic the rate and amount of leaf mass removed by one tobacco hornworm per plant. Undamaged leaves on these plants undergo a dramatic (457% for leaf position 5, 410% for leaf position 8) increase in total leaf alkaloids compared to same-age and positioned control leaves on undamaged control plants. However, potted greenhouse-grown plants fail to exhibit the same damage-induced increase in alkaloid content. The greenhouse environment differs from the field environment in factors known to influence leaf alkaloid content, particularly soil N, P, K, near-UV radiation, and relative humidity. However, altering these environmental factors does not make potted plants able to increase their leaf alkaloid levels in response to defoliation. Transplanting plants into larger pots with more soil does allow the plants to respond to defoliation. Thirty days after transplanting, the plants are again unresponsive to damage, probably as a result of becoming pot-bound. This result suggests a mechanism for the induction response, specifically that leaf damage triggers synthesis of these alkaloids in the roots, and offers a potentially valuable experimental tool for the study of induced-plant defenses in tobacco and other plants that synthesize alkaloids in their root tissues.     

The Dynamics of Carbon–Nutrient Balance: Effects of Cottonwood Acclimation to Short- and Long-Term Shade on Beetle Feeding Preferences

The carbon–nutrient balance hypothesis (CNBH) predicts that shading should increase leaf palatability to herbivores by decreasing concentrations of carbon (C) -based chemical defenses and increasing nitrogen (N). We measured cottonwood (Populus deltoids) growth, leaf chemistry, and beetle (Plagiodera versicolora) feeding preferences on saplings grown in either continuous high (HH) or low (LL) light, and saplings switched from high to low (HL) or low to high (LH) light for nine days. As expected, based on the CNBH, shading increased total N and decreased total phenol glycoside (C-based secondary metabolites) concentrations in plants from all shade treatments (LL, HL, and LH), relative to HH plants, with plant growth and gross leaf chemistry being affected by initial and final light regime. In contrast, while specific phenol glycoside concentrations were affected by the initial and final light regime, they also showed an initial × final light interaction. Beetles tended to prefer LL to HH plants. Beetles unexpectedly preferred HH to either HL or LH switched plants, most likely because high concentrations of a specific phenol glycoside – salicin – occurred in both switched treatments and inhibited beetle feeding. Plant chemical allocation during light acclimation led to unpredictable changes in specific C-based compounds, even though plant growth and gross chemistry conformed to expectations for shading effects and the CNBH. The response of this herbivore to altered concentrations of a specific compound confounded predictions based on average dynamics of suites of chemicals. Our findings may help explain why relationships between light availability and herbivory in field studies, where light varies on many time scales, can differ from those predicted by the CNBH. Understanding both dynamic plant chemical responses to altered resource availability and controls over allocation to specific compounds would likely enhance future predictability of specific environment-plant-herbivore interactions.     

Nitrogen competition in contour hedgerow systems in subtropical China

Contour hedgerow agroforestry has been studied for soil erosion control and soil fertility improvement in subtropical China. However, below-ground competitive and complementary interactions between tree hedges and crops have received relatively little attention in the scientific literature. A field experiment was conducted to explore the effects of a leguminous shrub hedge, false indigo (Amorpha fruticosa) and a non-legume gramineous hedge, vetiver (Vetiveria zizanioides), on the growth of soybean (Glycine max). Pot experiments were also carried out to determine the effect of below-ground interactions on nitrogen uptake between two contour hedgerow agroforestry with a ¹⁵N isotope method and root partition, i.e., a sheet barrier, a mesh barrier and no barrier. The results showed that the relative disadvantage of intercropping, expressed as land equivalent ratio, were 0.96/0.97 for the A. fruticosa–soybean system and 0.99 for the vetiver–soybean system, based on the dry matter (DM) production and N acquisition. Both area-adjusted yield and N content of soybean were significantly decreased in two intercropping treatments compared to those in the sole soybean treatment. The DM production of soybean, for example, was decreased by 10% and 5% under A. fruticosa and vetiver, respectively, when compared to the sole soybean. The intercropping disadvantage was mainly due to interspecific competitive interaction. The result was proved by lower yields and biomasses adjacent to the hedgerows. The ¹⁵N-based estimates of N uptake, in the vetiver–soybean system, of soybean with mesh separation (6.11 mg pot⁻¹) was lower than that (13.85 mg pot⁻¹) with no root separation, for vetiver the higher ¹⁵N uptake was observed in no root separation (13.90 mg pot⁻¹). In the A. fruticosa–soybean system, the lower ¹⁵N uptake of soybean (1.53 mg pot⁻¹) and A. fruticosa (6.42 mg pot⁻¹) were observed in no root separation. It is concluded that the growth of soybean was unexpectedly suppressed in two intercropping systems. The growth of A. fruticosa was clearly suppressed due to below-ground interactions, yet the growth of vetiver was improved to a great extent.     

Shade tree effects in an 8-year-old cocoa agroforestry system: biomass and nutrient diagnosis of <Emphasis Type="Italic">Theobroma cacao</Emphasis> by vector analysis

Farm product diversification, shade provision and low access to fertilizers often result in the purposeful integration of upper canopy trees in cocoa (Theobroma cacao) plantations. Subsequent modification to light and soil conditions presumably affects nutrient availability and cocoa tree nutrition. However, the level of complementarity between species requires investigation to minimize interspecific competition and improve resource availability. We hypothesized beneficial effects of upper canopy trees on cocoa biomass, light regulation, soil fertility and nutrient uptake. We measured cocoa standing biomass and soil nutrient stocks under no shade (monoculture) and under three structurally and functionally distinct shade trees: Albizia zygia (D.C.) Macbr, a nitrogen fixer; Milicia excelsa (Welw.), a native timber species; and Newbouldia laevis (Seem.), a native small stature species. Vector analysis was employed to diagnosis tree nutrition. Cocoa biomass was higher under shade (22.8 for sole cocoa versus 41.1 Mg ha⁻¹ for cocoa under Milicia), and declined along a spatial gradient from the shade tree (P < 0.05). Percent canopy openness differed between the three shade species (P = 0.0136), although light infiltration was within the optimal range for cocoa production under all three species. Soil exchangeable K was increased under Newbouldia, while available P decreased and total N status was unaffected under all shade treatments. Nutrient uptake by cocoa increased under shade (43–80% and 22–45% for N and P, respectively), with K (96–140%) as the most responsive nutrient in these multistrata systems. Addition of low-density shade trees positively affected cocoa biomass close to the shade tree, however proper management of upper stratum trees is required for optimum cocoa productivity and sustainability.     

Integrating legumes to improve N cycling on smallholder farms in sub-humid Zimbabwe: resource quality, biophysical and environmental limitations

The release of mineral-N in soil from plant residues is regulated by their ‘quality’ or chemical composition. Legume materials used by farmers in southern Africa are often in the form of litter with N concentration <2%. We investigated the decomposition of Sesbania sesban and Acacia angustissima litter in the field using litterbags, and N mineralization of a range of legume materials using a leaching tube incubation method in the laboratory. The mass loss of the litter could be described using a modified exponential decay model: Y = (Y ₀−Q)e^−kt + Q. The relative decomposition constants for Sesbania and Acacia litter were 0.053 and 0.039 d⁻¹, respectively. The % N mineralized from fresh Sesbania prunings was 55% compared with only 27% for the Sesbania litter after 120 days of incubation under leaching conditions. During the same period, fresh prunings of Acacia released only 12% of the added N while Acacia litter released 9%. Despite the large differences in N concentration between Acacia prunings and its litter, the total mineralized N was similar, as mineralization from prunings was depressed by the highly active polyphenols. While N supply may be poor, these slow decomposing litter materials are potentially useful for maintaining soil organic matter in smallholder farms. In two field experiments with contrasting soil texture, Sesbania, Acacia and Cajanus produced large amounts of biomass (>5 Mg ha⁻¹) and improved N cycling significantly (>150 kg N ha⁻¹) on the clay loam soil, but adapted poorly on the sandier soil. There was a rapid N accumulation in the topsoil at the beginning of the rains in plots where large amounts of Sesbania or Acacia biomass had been incorporated. Despite the wide differences in resource quality between these two, there was virtually no difference in N availability in the field as this was, among other factors, confounded by the quantity of N added. A substantial amount of the nitrate was leached to greater than 0.4 m depth within a three-week period. Also, the incidence of pests in the first season, and drought in the second season resulted in poor nitrogen use efficiency. Our measurements of gaseous N losses in the field confirmed that N₂O emissions were <0.5 kg N ha⁻¹. As we had measurements of all major N flows, we were able to construct overall N budgets for the improved fallow – maize rotation systems. These budgets indicated that, in a normal rainfall season with no major pest problems, reducing nitrate leaching would be the single largest challenge to increased N recovery of added organic N in the light textured soils.     

Phytotoxicity of <Emphasis Type="Italic">Phytolacca americana</Emphasis> Leaf Extracts on the Growth,and Physiological Response of <Emphasis Type="Italic">Cassia mimosoides</Emphasis>

We examined the allelochemical effects of control soil, native soil (treated soil), and leaf extracts of Phytolacca americana (pokeweed) on the germination rate and seedling growth of Cassia mimosoides var. nomame. We also studied the resulting changes in root-tip ultrastructure and peroxidase isozyme biochemistry. P. americana leaf extract inhibited seed germination, seedling growth, and biomass when compared to control and treated soil. Root and shoot growth in treated soil was stimulated relative to control soil, but root growth was inhibited by 50% in the leaf extract treatment. Biomass of C. mimosoides seedlings grown on leaf extract was reduced sevenfold when compared to the control seedlings. The amounts of total phenolic compounds in the leaf extract, treated soil, and control soil were 0.77, 0.14, and 0.03 mg l⁻¹, respectively. The root tips of C. mimsoides treated with leaf extracts of P. americana showed amyloplasts and large central vacuoles with electron-dense deposits inside them when compared to control root tips. The activity of guaiacol peroxidase (GuPOX) in whole plant, roots, and shoots of C. mimosoides increased as leaf extract increased; maximum activity was observed in extract concentrations of 75% and higher. Root GuPOX activity was three times higher than in shoots. Therefore, we conclude that inhibition of C. mimosoides growth is related to the phenolic compounds in the P. americana leaf extract and the ultrastructure changes in root-tip cells and increased GuPOX activity is a response to these allelochemicals.     

Allelopathic research of subtropical vegetation in Taiwan II. Comparative exclusion of understory byPhyllostachys edulis andCryptomeria japonica

On many hillsides of Taiwan there is a unique pattern of weed exclusion byPhyllostachys edulis (bamboo) andCryptomeria japonica (conifer) in which the density, diversity, and dominance of understory species are very different. Although the physical conditions of light, soil moisture, and soil nutrients strongly favor the growth of understory in a bamboo community, the biomass of its undergrowth is significantly low, indicating that physical competition among the understory species in the bamboo and conifer communities does not cause the observed differences. However, the biochemical inhibition revealed by these two plants appeared to be an important factor. The growth ofPellionia scabra seedlings, transplanted from the study site into greenhouse pots, was evidently suppressed by the aqueous leachate of bamboo leaves but was stimulated by that of conifer leaves. The radicle growth of lettuce, rye grass, and rice plants was also clearly inhibited by the leachate and aqueous extracts of bamboo leaves but not by those of conifer leaves. Six phytotoxins,o-hydroxyphenylacetic,p-hydroxybenzoic,p-coumaric, vanillic, ferulic, and syringic acids were found in the aqueous leachate and extracts of leaves and alcoholic soil extracts ofP. edulis, while the first three compounds were absent in the extracts ofC. japonica. The phytotoxicities of extracts were correlated with the phytotoxins present in both leaves and soils. The understory species might be variously tolerant to the allelopathic compounds produced by the two plants, resulting in a differential selection of species underneath. Therefore, comparative allelopathic effects ofPhyllostachys edulis andCryptomeria japonica may play significant roles in regulating the populations of the understories.Paper No. 253 of the Scientific Journal Series of the Institute of Botany, Academia Sinica, Taipei, Taiwan. This study was supported by the National Science Council of the Republic of China.     

Evaluation of allelopathic potential of dominant herbaceous species in a coffee plantation

Several shaded coffee plantations in Coatepec, Veracruz (Mexico) are characterized by a dense cover of herbaceous vegetation mainly dominated by species from the Commelinaceae which protect the soil from erosion and presumably contribute to regulating the abundance of other weeds. To detect their alleopathic potential, leachates from fresh, air-dried, or oven-dried plants and litter collected during different months of the year were tested uponBrassica campestris, Bidens pilosa, andRumex sp. seeds. Significant radicle growth inhibitions were obtained mainly from dried plants and litter collected during the rainy season (August). Drainage water collected from pots with fresh, chopped plants and litter produced no inhibitions until the third week of recycling the water. Concentrated soil extracts from chopped plants and litter collected after seven weeks of decomposition produced significant inhibitions on radicle growth ofRumex sp. Dry weight ofBidens pilosa was significantly reduced when grown in soils treated with fresh and chopped plants and litter exposed to natural field conditions for five weeks.