Nitrogen fertilisation reduces sink strength of poplar ectomycorrhizae during recovery after drought more than phosphorus fertilisation

Plant and Soil - Low nitrogen (N) use efficiency endangers world crop production and ground water. Biochar has been well documented in sequestering C in agricultural soils and improving soil...     

Nitrogen fertilisation and light effects on growth and photosynthesis of lodgepole pine seedlings

Seedlings of Pinus contorta were grown for 16 wk without ectomycorrhizas in the greenhouse at three levels of irradiance (100, 210 and 470 μEm^-2s^-1) and three levels of ammonium nitrate (3, 62 and 248 mg litre^-1 of N). Measurements at 5, 10 and 16 wk of age indicated that plants receiving the highest levels of irradiance and nitrogen had significantly larger biomass, a higher total foliar N and P content and net photosynthesis per unit leaf area than those at lower levels of each factor. Although the root/shoot ratio also increased from low to high irradiance at each harvest, nitrogen application resulted in an increased ratio from 3 to 62 mg litre^-1 N, but decreased ratio at 248 mg litre^-1 N.     

Responses of Fraxinus excelsior seedlings to grass-induced above- and below-ground competition

The competitive interactions between woody seedlings and herbaceous vegetation have received increasing interest in recent years. However, little is known about the relative contributions and underlying mechanisms of above- and below-ground competition between species. We used a novel experimental approach to assess the responses of Fraxinus excelsior seedlings to different combinations of root and shoot competition imposed by the grass Dactylis glomerata under greenhouse conditions. Seedling growth was significantly reduced by competition for soil resources, but neither biomass nor height were significantly affected by shoot competition for light. Competitive response indices based on biomass confirmed that below-ground competition was more important than above-ground competition, and indicated that root and shoot competition did not interact to influence plant growth. Fraxinus biomass allocation and seedling traits were almost all significantly affected by root competition; these responses varied depending on the trait examined. In contrast, morphological responses to shoot competition were limited. In the absence of root competition, seedlings showed a significant increase in the biomass allocated to leaves and a greater leaf area ratio in response to shoot competition. Our findings suggest that morphological modifications help to mitigate the negative effects of competition, but the expression of plasticity may be suboptimal due to resource constraints. The present study also highlights the importance of appropriate experimental controls and analysis to avoid confounding effects of experimental design and ontogeny on the interpretation of competitive responses.     

Nitrogenase activity and N2 fixation are stimulated by elevated CO2 in a tropical N2-fixing tree

 Seeds of Gliricidia sepium, a fast-growing woody legume native to seasonal tropical forests of Central America, were inoculated with N₂-fixing Rhizobium bacteria and grown in environmentally controlled glasshouses for 67–71 days under ambient CO₂ (35 Pa) and elevated CO₂ (70 Pa) conditions. Seedlings were watered with an N-free, but otherwise complete, nutrient solution such that bacterial N₂ fixation was the only source of N available to the plant. The primary objective of our study was to quantify the effect of CO₂ enrichment on the kinetics of photosynthate transport to nodules and determine its subsequent effect on N₂ fixation. Photosynthetic rates and carbon storage in leaves were higher in elevated CO₂ plants indicating that more carbon was available for transport to nodules. A ¹⁴CO₂ pulse-chase experiment demonstrated that photosynthetically fixed carbon was supplied by leaves to nodules at a faster rate when plants were grown in elevated CO₂. Greater rates of carbon supply to nodules did not affect nodule mass per plant, but did increase specific nitrogenase activity (SNA) and total nitrogenase activity (TNA) resulting in greater N₂ fixation. In fact, a 23% increase in the rate of carbon supplied to nodules coincided with a 23% increase in SNA for plants grown in elevated CO₂, suggesting a direct correlation between carbon supply and nitrogenase activity. The improvement in plant N status produced much larger plants when grown in elevated CO₂. These results suggest that Gliricidia, and possibly other N₂-fixing trees, may show an early and positive growth response to elevated CO₂, even in severely N-deficient soils, due to increased nitrogenase activity. Received: 27 February 1996 / Accepted: 19 June 1996     

Animal treading during wet soil conditions reduces N2 fixation in mixed clover-grass pasture

A field experiment was carried out over 12 months to determine the effect of animal treading on N₂ fixation in a mixed white clover-ryegrass pasture. The experimental site was defoliated by mowing for the duration of the study. A single treading event of moderate or severe pugging intensity was initiated in plots during wet spring conditions by using dairy cows at varying stocking rates (4.5 cows 100 m⁻² for 1.5 or 2.5 h, respectively). Inputs of dung and urine onto the plots was avoided by overnight housing of the cows and interception of excreta during the pugging event. Soil air-filled porosity decreased from 21% in the non-pugged control to 15–16% in pugged treatments by day 3. Bulk density of soil was not significantly affected by pugging. Soil inorganic N concentration increased in pugged treatments, and was 4-fold greater on day 28 in severely pugged plots compared to non-pugged plots. White clover plant density and plant size was markedly lower in pugged treatments (up to 85% and 72% reduction, respectively under severe pugging). White clover growth was most affected during the first 156 days after pugging (up to 90% decrease under severe pugging), leading to an annual clover dry matter production loss of 9% and 52%, respectively. The proportion of clover N derived from atmospheric N₂ (%Ndfa; estimated by ¹⁵N dilution) was initially reduced (to a lower limit of 43%) by severe pugging (days 28–71) before recovery to control levels (90%) by day 91. Annual N₂ fixation in clover herbage decreased significantly from 76 kg N ha⁻¹ yr⁻¹ in the non-pugged control, to 66 and 36 kg N ha⁻¹ yr⁻¹ under moderate and severe pugging, respectively. Most of this difference was evident within the first 156 days after pugging. Our data indicates that the major loss in fixed N₂ input under pugging was due to reduced clover growth and production resulting from pugging damage and loss of residual white clover biomass by hoof action.     

A continental-scale analysis of tree cover in African savannas

Aim We present a continental‐scale analysis that explores the processes controlling woody community structure in tropical savannas. We analyse how biotic and abiotic factors interact to promote and modify tree cover, examine alternative ecological hypotheses and quantify disturbance effects using satellite estimates of tree cover. Location African savannas. Methods Tree cover is represented as a resource‐driven potential cover related to rainfall and soil characteristics perturbed by natural and human factors such as fire, cattle grazing, human population and cultivation. Within this framework our approach combines semi‐empirical modelling and information theory to identify the best models. Results Woody community structure across African savannas is best represented by a sigmoidal response of tree cover to mean annual precipitation (MAP), with a dependency on soil texture, which is modified by the separate effects of fire, domestic livestock, human population density and cultivation intensity. This model explains c. 66% of the variance in tree cover and appears consistent across the savanna regions of Africa. Main conclusions The analysis provides a new understanding of the importance and interaction of environmental and disturbance factors that create the broad spatial patterns of tree cover observed in African savannas. Woody cover increases with rainfall, but is modified by disturbances. These ‘perturbation’ effects depend on MAP regimes: in arid savannas (MAP < 400 mm) they are generally small (< 1% decrease in cover), while in semi‐arid and mesic savannas (400–1600 mm), perturbations result in an average 2% (400 mm) to 23% (1600 mm) decrease in cover; fire frequency and human population have more influence than cattle, and cultivation appears, on average, to lead to small increases in woody cover. Wet savannas (1600–2200 mm) are controlled by perturbations that inhibit canopy closure and reduce tree cover by, on average, 24–34%. Full understanding of the processes determining savanna structure requires consideration of resource limitation and disturbance dynamics.     

Large herbivores may alter vegetation structure of semi-arid savannas through soil nutrient mediation

In savannas, the tree-grass balance is governed by water, nutrients, fire and herbivory, and their interactions. We studied the hypothesis that herbivores indirectly affect vegetation structure by changing the availability of soil nutrients, which, in turn, alters the competition between trees and grasses. Nine abandoned livestock holding-pen areas (kraals), enriched by dung and urine, were contrasted with nearby control sites in a semi-arid savanna. About 40 years after abandonment, kraal sites still showed high soil concentrations of inorganic N, extractable P, K, Ca and Mg compared to controls. Kraals also had a high plant production potential and offered high quality forage. The intense grazing and high herbivore dung and urine deposition rates in kraals fit the accelerated nutrient cycling model described for fertile systems elsewhere. Data of a concurrent experiment also showed that bush-cleared patches resulted in an increase in impala dung deposition, probably because impala preferred open sites to avoid predation. Kraal sites had very low tree densities compared to control sites, thus the high impala dung deposition rates here may be in part driven by the open structure of kraal sites, which may explain the persistence of nutrients in kraals. Experiments indicated that tree seedlings were increasingly constrained when competing with grasses under fertile conditions, which might explain the low tree recruitment observed in kraals. In conclusion, large herbivores may indirectly keep existing nutrient hotspots such as abandoned kraals structurally open by maintaining a high local soil fertility, which, in turn, constrains woody recruitment in a negative feedback loop. The maintenance of nutrient hotspots such as abandoned kraals by herbivores contributes to the structural heterogeneity of nutrient-poor savanna vegetation.     

Seasonal physiological responses of Argania spinosa tree from Mediterranean to semi-arid climate

Argania spinosa (the argan tree) is a slow-growing tree endemic of Morocco, growing on semi-arid areas where no other tree species can live. With the aim of predicting temporal changes in A. spinosa woodlands under a probable increase in aridity, we set off to investigate these questions: how do A. spinosa physiological attributes respond to variations in climatic conditions and seasonality, and which is the set of attributes that most affects tree response to environmental conditions? In three study sites, Beni Snassen (North), High-Atlas (Mountain) and Admine Forest in Agadir (Coastal), gas exchange measurements, photochemical efficiency, leaf water potential and different leaf attributes were monitored in February, July and November of 2006. The Mountain site presents the most continental climate. Trees in this site were the most stressed in summer, having the lowest midday leaf water potential values, photochemical efficiency and assimilation rates. We found a Ψ_md threshold around -4 MPa, below which stomatal conductance responds linearly to Ψ_md. Plants from the North area never reached this threshold during the study period. Although leaf pigments presented a clear seasonal pattern, leaves from Coastal trees exhibit the highest content for each season. The three study sites were separated by two discriminate functions obtained by canonical discriminant analysis. In summer, the Mountain population is separated from the other sites mainly by assimilation rate and F_v/F_m, while in winter transpiration rates and chlorophyll content are the main discriminant variables. Our study shows that A. spinosa trees adjust their physiological status and leaf attributes to environmental conditions allowing plants to thrive under a dry climate. Under a scenario of global change, the distribution of the argan tree likely shifts to milder areas.     

Assessing the symbiotic dependency of grain and tree legumes on N2 fixation for their N nutrition in five agro-ecological zones of Botswana

To assess the symbiotic dependency of grain and shrub/tree legumes within five agro-ecological zones of Botswana, fully expanded leaves of the test species were sampled from about 26 study sites within Ngwaketse, Gaborone, Central, Ghanzi and Kalahari agro-ecological zones. Isotopic analysis revealed significant differences in δ¹⁵N values of the grain legumes [cowpea (Vigna unguiculata L. Walp), Bambara groundnut (Vigna subterranea L. Verde.), and groundnut (Arachis hypogaea L.)] from the 26 farming areas in both 2005 and 2006. Estimates of %Ndfa of leaves also showed significant differences between farming areas, with cowpea deriving more than 50% of its N nutrition from symbiotic fixation. In terms of distribution, many more symbiotic shrub/tree species were found in the wetter Ngwaketse agro-zone compared to the fewer numbers in the drier Kalahari region. Acacias were the more dominant species at all sites. Leaf δ¹⁵N values of shrub/tree species also varied strongly across Botswana, with 11 out of 18 of these legumes deriving about 50%, or more, of their N from symbiotic N₂ fixation.Acacia caffra, in particular, obtained as much as 93.6% of its N nutrition from symbiotic fixation in the wetter Ngwaketse agro-zone. This study has shown that grain legumes sampled from farmer’s fields in Botswana obtained considerable amounts of their N from symbiotic fixation. We have also shown that shrub and tree legumes probably play an important role in the N economy of the savanna ecosystems in Botswana. However, the decline in the number of functional N₂-fixing shrub/tree legumes along an aridity gradient suggests that soil moisture is a major constraint to N₂ fixation in the tree legumes of Botswana.     

Current range condition in relation to land management systems in semi-arid savannas of Swaziland

Rangeland condition was assessed in the lowveld of Swaziland to determine the current status with emphasis on contrasting different land management systems and soil classes in two study areas. The assessment incorporated the grass and woody plant layer. The methods employed to evaluate the grass layer were ecological condition index (ECI) and weighted palatability composition (WPC). The government ranch had significantly higher ECI (mean 714.5) and WPC (mean 61) than the communal land (mean: ECI – 533.5; WPC – 48.7) and the game reserve (mean: ECI – 578.9; WPC – 47.9). The ECI and WPC values did not differ between the soil classes in most cases (range: ECI 551.5–645.9 and WPC – 43.7–57.6). The density of all woody plants and encroaching plants alone were the highest in communal land. Basal cover ranged from poor (2–3%) to good (>5%), while bare ground was rated from low (<1%) to high (>5%). Overall results showed great variability of studied variables at site and landscape levels of resolution. Generally, the grass layer was rated between fair and good when assessed on the basis of ecological and palatability merits. There was a clear indication of advancement of bush encroachment.     

Nitrogen fixation and methanotrophy in forest mosses along a N deposition gradient

Nitrogen deposition has decreased the plant-associated nitrogen (N₂) fixation when measured using the indirect acetylene reduction assay (ARA). However, nitrogen deposition can also lead to changes in the diversity of moss symbionts, e.g. affect methanotrophic N₂ fixation, which is not measured by ARA. To test this hypothesis we compared ARA with the direct stable isotope method (¹⁵N₂ incorporation) and studied methanotrophy in two mosses, Hylocomium splendens and Pleurozium schreberi, collected from seven forest sites along a boreal latitudinal N deposition transect. We recognized that the two independent N₂ fixation measures gave corresponding results with the conversion factor of 3.3, but the ¹⁵N₂ method was more sensitive for finding a signal of low N₂ fixation activity. Methane carbon fixation associated with mosses was under the detection limit (<2 nmol C g⁻¹ h⁻¹). N₂ fixation rates were more pronounced in the mosses with higher C/N ratio, and in the green upper parts of the shoot than in the lower brownish parts. Sequencing of nifH genes revealed that dominating diazotrophs were affiliated to cyanobacterial genera Nostoc and Nodularia, but methanotrophic diazotrophs were not found in the nifH libraries. We conclude that the suppression of N₂ fixation along the deposition gradient was consistent regardless of the measurement technique, and microbial community changes toward methanotrophic or otherwise acetylene-sensitive N₂ fixation could not explain this trend.     

Influence of urea on biological N2 fixation and N transfer from Azolla intercropped with rice

The N₂ fixed by Azolla before and after urea application during the rice cycle, the mineralisation of Azolla-N as well as its availability to rice was studied in two greenhouse experiments conducted in 1996 and 1997 and in June 1998 in Goettingen (Germany). Dry matter production of the various rice parts of experiment 1 showed a clear positive synergism between treatment with Azolla and urea with a resulting apparent N recovery by rice increasing from 40% (without Azolla) to 57% in the presence of Azolla. Part of this increase may be due to N fixed biologically by Azolla and transferred to the rice. The second experiment shed some light on the role of BNF. Using an iterative method of estimation, the daily rate of N fixation was estimated at 0.6 – 0.7 kg N ha^–1. The rate was not so much affected by the age of the Azolla crop. At this rate, the BNF would amount to up to 100 kg N ha^–1 over a 130-day season. Assuming that BNF may be inhibited for a period of 5 – 10 days following urea application due to high levels of N in the floodwater, this might reduce the BNF by between 6 and 14 kg N ha over the season. Using the mean-pool-abundance concept, it was estimated that around 75 – 80% of the Azolla-N mineralized during the growth period was actually absorbed by the rice plants. Of the N taken up by rice around 28% was derived from the biologically fixed Azolla N, the remainder was urea N cycled through the Azolla. Azolla also seems to help sustain the soil N supply by returning N to the soil in quantities roughly equal to those extracted from the soil by the rice plant.     

A comparison between measured and modeled N2O emissions from Inner Mongolian semi-arid grassland

The objectives of this study were (1) to determine the effect of land use on N₂O emissions from Inner Mongolian semi-arid grasslands of China and (2) to evaluate the process-based DNDC model to extrapolate our field measurements from a limited number of sites to a larger temporal and spatial scale. The results suggest the following. Rainfall event was the dominant controlling factor for the seasonal variations of the N₂O fluxes. The seven selected sites exhibited a similar seasonal trend in N₂O emission, despite their different vegetation, land use and textures. In the typical steppe, N₂O fluxes generally decrease with decreasing soil organic C (SOC) and total N content, indicating that soil C and N pools are very important in determining the spatial magnitude of the N₂O flux. N₂O emissions were very small during the entire growing season, averaging only 0.76 g N₂O-N ha^–1 day^–1 for the five typical steppe sites, 0.35 g N₂O-N ha^–1 day^–1 for the mown meadow steppe site, and 0.83 g N₂O-N ha^–1 day^–1 from the cropped meadow steppe site. No enhanced effect due to overgrazing was observed for the N₂O emission from the semi-arid grasslands. This was mainly results from the decreased SOC content due to overgrazing, which may have reduced the promoting effect of increased soil bulk density by trampling and animal excreta. Except for the mown steppe site, the model predictions of the N₂O flux for the six different sites agree well with the observed values (r ² ranging from 0.35 to 0.68). It would be concluded that the DNDC model captured the key driving process for N₂O emission. Nitrification was the predominant process, contributing 64–88% to the N₂O emission. However, in terms of the magnitude of the N₂O emission, further modifications should focus on the underestimated N₂O flux during the spring and autumn periods (nitrification, freeze/thaw cycles) and the effect of topography and the mowing on N₂O emission.     

Sudden exposure to solar UV-B radiation reduces net CO(2) uptake and photosystem I efficiency in shade-acclimated tropical tree seedlings

Tree seedlings developing in the understory of the tropical forest have to endure short periods of high-light stress when tree-fall gaps are formed, and direct solar radiation, including substantial UV light, reaches the leaves. In experiments simulating the opening of a tree-fall gap, the response of photosynthesis in leaves of shade-acclimated seedlings (Anacardium excelsum, Virola surinamensis, and Calophyllum longifolium) to exposure to direct sunlight (for 20-50 min) was investigated in Panama (9 degrees N). To assess the effects of solar UV-B radiation (280-320 nm), the sunlight was filtered through plastic films that selectively absorbed UV-B or transmitted the complete spectrum. The results document a strong inhibition of CO(2) assimilation by sun exposure. Light-limited and light-saturated rates of photosynthetic CO(2) uptake by the leaves were affected, which apparently occurred independently of a simultaneous inhibition of potential photosystem (PS) II efficiency. The ambient UV-B light substantially contributed to these effects. The photochemical capacity of PSI, measured as absorbance change at 810 nm in saturating far-red light, was not significantly affected by sun exposure of the seedlings. However, a decrease in the efficiency of P700 photooxidation by far-red light was observed, which was strongly promoted by solar UV-B radiation. The decrease in PSI efficiency may result from enhanced charge recombination in the reaction center, which might represent an incipient inactivation of PSI, but contributes to thermal dissipation of excessive light energy and thereby to photoprotection.     

Effects of NaCl on growth, water status, N2 fixation, and ion distribution in Pterocarpus officinalis seedlings

Pterocarpus officinalis (Fabaceae) dominates in the swamp forests of the Lesser Antilles, submitted to strong variations of soil salinity (30–445 mM). This study aimed to assess the effect of salinity on growth, nodulation, N₂ fixation, water status and ions content in P. officinalis and to clarify the mechanisms involved. Seedlings inoculated or not with two strains from areas of contrasting salinity levels (< to 50 or 445 mM) were watered with 0, 171 and 342 mM solutions of NaCl in greenhouse conditions. Non-inoculated seedlings were tolerant to a salinity of 171 mM, with no significant effect on seedling biomass. Evapotranspiration per unit of leaf area (E/TLa) remained unchanged at 171 mM. Maintenance of a constant E/TLa and especially the control of ion transport to the upper parts of the plant could explain seedling salt tolerance up to intermediate salinity conditions (171 mM). The two strains have a 99.8% genetic identity in spite of differences in their original habitats, this explaining the similar response of the symbiosis to salinity. The higher salt sensitivity of inoculated seedlings was linked to the sensitivity of both Bradyrhizobium strains (reduction of free-living cells) and to that of the nodulation process (fewer nodules and inhibition of N₂-fixation) to intermediate salinity.     

Performance of seedlings of the invasive alien tree Schinus molle L. under indigenous and alien host trees in semi-arid savanna

We assessed the importance of host trees in influencing invasion patterns of the alien tree Schinus molle L. ( Anacardiaceae ) in semi-arid savanna in South Africa. Recruitment of S. molle is dependent on trees in its invaded habitat, particularly Acacia tortilis Hayne. Another leguminous tree, the invasive alien mesquite ( Prosopis sp.), has become common in the area recently, but S. molle rarely recruits under canopies of this species. Understanding of the association between these species is needed to predict invasion dynamics in the region. We conducted experiments to test whether: (i) seedling survival of S. molle is better beneath A. tortilis than beneath mesquite canopies; (ii) growth rates of S. molle seedlings are higher beneath A. tortilis than beneath mesquite. Results showed that growth and survival of S. molle did not differ significantly beneath the native A. tortilis and the alien Prosopis species. This suggests that microsites provided by canopies of mesquite are as good for S. molle establishment as those provided by the native acacia. Other factors, such as the failure of propagules to arrive beneath mesquite trees, must be sought to explain the lack of recruitment beneath mesquite.     

Nitrogen fixation by Spirillum sp. from rice roots

Nitrogen fixation by Spirillum sp. obtained from the roots of rice plants grown at different levels of combined nitrogen was studied. The roots of rice plants exposed to low levels of combined nitrogen (20–40 kg N/ha) harboured Spirillum sp. possessing higher nitrogen-fixing efficiency as compared to the cultures from plants receiving 60–100 kg N/ha. More-over, the nitrogen-fixing efficiency of these Spirillum spp. varied with age of the plant, irrespective of the dosage of combined nitrogen.     

Nitrogen fixation (15N dilution) with soybeans under Thai field conditions

Controlled environment and field studies were conducted to determine relationships between various measurements of N₂ fixation using soybeans and to use these measures to evaluate a number ofBradyrhizobium japonicum strains for effectiveness in N₂ fixation in Thai soils.¹⁵N dilution measurements of N₂ fixation showed levels of fixation ranging from 32 to 161 kg N ha⁻¹ depending on bacterial strain, host cultivar and location. Midseason measures of N₂ fixation were correlated with each other, but not related measures taken at maturity. Ranking ofB. japonicum strains based on performance under controlled conditions in N-free media were highly correlated with rankings based on soybean seed yields and N₂ fixation under field conditions. This study showed that inoculation of soybeans with effectiveB. japonicum strains can result in significant increases in yield and uptake of N through fixation. The most effective strains tested for use in Thai conditions were those isolated from Thai soils; however, effective strains from other locations were also of benefit.