Per Ambus

Interspecific competition, N use and interference with weeds in pea–barley intercropping

Field pea (Pisum sativum L.) and spring barley (Hordeum vulgare L) were inter- and sole cropped to compare the effects of crop diversity on productivity and use of N sources on a soil with a high weed pressure. 15N enrichment techniques were used to determine the pea-barley-weed-N dynamics. The pea-barley intercrop yielded 4.6 t grain ha-1, which was significantly greater than the yields of pea and barley in sole cropping. Calculation of Land Equivalent Ratios showed that plant growth factors were used from 25 to 38% more efficiently by the intercrop than by the sole crops. Barley sole crops accumulated 65 kg soil N ha-1 in aboveground plant parts, which was similar to 73 kg soil N ha-1 in the pea-barley intercrop and significantly greater than 15 kg soil N ha-1 in the pea sole crop. The weeds accumulated 57 kg soil N ha-1 in aboveground plant parts during the growing season in the pea sole crops. Intercropped barley accumulated 71 kg N ha-1. Pea had to rely on N2 fixation with 90-95% of aboveground N accumulation being derived from N2 fixation independent of cropping system. Pea grown in intercrop with barley instead of sole crop had a better competitive ability towards weeds and soil inorganic N was consequently used for barley grain production instead of weed biomass. There was no indication of a greater inorganic N content after pea compared to barley or pea-barley. However, 46 days after emergence there was about 30 kg N ha-1 inorganic N more under the pea sole crop than under the other two crops. Such greater inorganic N levels during early growth phases was assumed to induce aggressive weed populations and interspecific competition. Pea-barley intercropping seems to be a promising practise of protein production in cropping system with high weed pressures and low levels of available N.

The comparison of nitrogen use and leaching in sole cropped versus intercropped pea and barley

The effect of sole and intercropping of field pea (Pisumsativum L.) and spring barley (Hordeum vulgareL.) and of crop residue management on crop yield,NO3− leaching and N balance in the cropping systemwas tested in a 2-year lysimeter experiment on a temperate sandy loam soil. Thecrop rotation was pea and barley sole and intercrops followed by winter-rye anda fallow period. The Land Equivalent Ratio (LER), which is defined as therelative land area under sole crops that is required to produce the yieldsachieved in intercropping, was used to compare intercropping performancerelative to sole cropping. Crops received no fertilizer in the experimentalperiod. Natural 15N abundance techniques were used to determine peaN2 fixation. The pea–barley intercrop yielded 4.0 Mg grainha−1, which was about 0.5 Mg lowerthan theyields of sole cropped pea but about 1.5 Mg greater than harvestedin sole cropped barley. Calculation of the LER showed thatplant growth resources were used from 17 to 31% more efficiently by theintercrop than by the sole crops. Pea increased the N derived fromN2fixation from 70% when sole cropped to 99% of the total aboveground Naccumulation when intercropped. However, based upon aboveground N accumulationthe pea–barley intercrop yielded about 85 kg Nha−1, which was about 65 kg lower thansolecropped pea but about three times greater than harvested in sole croppedbarley.Despite different preceding crops and removal or incorporation of straw, therewas no significant difference between the subsequent non-fertilized winter-ryegrain yields averaging 2.8 Mg ha−1, indicating anequalization of the quality of incorporated residue by theNO3− leaching pattern.NO3− leaching throughout the experimental periodwas61 to 76 kg N ha−1. Leaching dynamics indicateddifferences in the temporal N mineralization comparing lysimeters previouslycropped with pea or with barley. The major part of this N was leached duringautumn and winter. Leaching tended to be smaller in the lysimeters originallycropped with the pea–barley intercrops, although not significantly differentfromthe sole cropped pea and barley lysimeters. Soil N balances indicated depletionof N in the soil–plant system during the experimental period, independent ofcropping system and residue management. N complementarity in the croppingsystemand the synchrony between residual N availability and crop N uptake isdiscussed.

Temporal and spatial distribution of roots and competition for nitrogen in pea-barley intercrops – a field study employing 32P technique

Root system dynamics, productivity and N use were studied in inter- and sole crops of field pea (Pisum sativum L.) and spring barley (Hordeum vulgare L.) on a temperate sandy loam. A 32P tracer placed at a depth of 12.5, 37.5, 62.5 or 87.5 cm was employed to determine root system dynamics by sampling crop leaves at 0, 15, 30 and 45 cm lateral distance. 15N addition was used to estimate N2 fixation by pea, using sole cropped barley as reference crop. The Land Equivalent Ratio (LER), which is defined as the relative land area under sole crops that is required to produce the yields achieved in intercropping, were used to compare the crop growth in intercrops relative to the respective sole crops.The 32P appearance in leaves revealed that the barley root system grows faster than that of pea. P uptake by the barley root system during early growth stages was approximately 10 days ahead of that of the pea root system in root depth and lateral root distribution. More than 90% of the P uptake by the pea root system was confined to the top 12.5 cm of soil, whereas barley had about 25–30% of tracer P uptake in the 12.5 – 62.5 cm soil layer. Judging from this P uptake, intercropping caused the barley root system to grow deeper and faster lateral root development of both species was observed. Barley accumulated similar amounts of aboveground N when grown as inter- and sole crop, whereas the total aboveground N acquired by pea in the intercrop was only 16% of that acquired in the pea sole crop. The percentage of total aboveground N derived from N2 fixation in sole cropped pea increased from 40% to 80% during the growth period, whereas it was almost constant at 85% in intercropped pea. The total amounts of N2 fixed were 95 and 15 kg N ha−1 in sole cropped and intercropped pea, respectively. Barley was the dominant component of the pea-barley intercrop, obtaining 90% of its sole crop yield, while pea produced only 15% of the grains of a sole crop pea. Intercropping of pea and barley improved the utilization of plant growth resources (LER > 1) as compared to sole crops. Root system distribution in time and space can partly explain interspecific competition. The 32P methodology proved to be a valuable tool for determining root dynamics in intercropping systems.

Enzymatic Evidence for the Key Role of Arginine in Nitrogen Translocation by Arbuscular Mycorrhizal Fungi

Key enzymes of the urea cycle and 15N-labeling patterns of arginine (Arg) were measured to elucidate the involvement of Arg in nitrogen translocation by arbuscular mycorrhizal (AM) fungi. Mycorrhiza was established between transformed carrot (Daucus carota) roots and Glomus intraradices in two-compartment petri dishes and three ammonium levels were supplied to the compartment containing the extraradical mycelium (ERM), but no roots. Time courses of specific enzyme activity were obtained for glutamine synthetase, argininosuccinate synthetase, arginase, and urease in the ERM and AM roots. 15NH4+ was used to follow the dynamics of nitrogen incorporation into and turnover of Arg. Both the absence of external nitrogen and the presence of l-norvaline, an inhibitor of Arg synthesis, prevented the synthesis of Arg in the ERM and resulted in decreased activity of arginase and urease in the AM root. The catabolic activity of the urea cycle in the roots therefore depends on Arg translocation from the ERM. 15N labeling of Arg in the ERM was very fast and analysis of its time course and isotopomer pattern allowed estimation of the translocation rate of Arg along the mycelium as 0.13 μg Arg mg−1 fresh weight h−1. The results highlight the synchronization of the spatially separated reactions involved in the anabolic and catabolic arms of the urea cycle. This synchronization is a prerequisite for Arg to be a key component in nitrogen translocation in the AM mycelium.

Biomass production, symbiotic nitrogen fixation and inorganic N use in dual and tri-component annual intercrops

The interspecific complementary and competitive interactions between pea (Pisum sativum L.), barley (Hordeum vulgare L.) and oilseed rape (Brassica napus L.), grown as dual and tri-component intercrops were assessed in a field study in Denmark. Total biomass production and N use at two levels of N fertilisation (0.5 and 4.0 g N/m2), were measured at five harvests throughout a growing season. All intercrops displayed land equivalent ratio values close to or exceeding unity, indicating complementary use of growth resources. Whereas both rape and barley responded positively to increased N fertilisation, irrespective of whether they were grown as sole- or intercrops, pea was strongly suppressed when grown in intercrop. Of the three crops barley was the strongest competitor for both soil and fertiliser N, rape intermediate and pea the weakest. Faster initial growth of barley than pea and rape gave barley an initial competitive advantage, an advantage that in the two dual intercrops was strengthened by the addition of N. Apparently the competitive superiority of barley was less strong in the tri-component intercrop, indicating that the impact of the dominantmay, through improved growth of both rape and pea, have been diminished through indirect facilitation. Interspecific competition had a promoting effect on the percent of nitrogen derived from N2 fixation of pea, and most so at the low N fertilisation level. Results indicate that the benefits achieved from the association of a legume and nonlegume, in terms of N2 fixed were greatest when pea was grown in association with rape as opposed to barley which could indicate that the benefits achieved from the association of a legume and nonlegume are partly lost if the nonlegume is too strong a competitor.

Nitrogen processes in terrestrial ecosystems

Executive summary Nature of the problem Nitrogen cycling in terrestrial ecosystems is complex and includes microbial processes such as mineralization, nitrification and denitrification, plant physiological processes (e.g. nitrogen uptake and assimilation) and physicochemical processes (leaching, volatilization). In order to understand the challenges nitrogen puts to the environment, a thorough understanding of all these processes is needed. Approaches This chapter provides an overview about processes relating to ecosystem nitrogen input and output and turnover. On the basis of examples and literature reviews, current knowledge on the effects of nitrogen on ecosystem functions is summarized, including plant and microbial processes, nitrate leaching and trace gas emissions. Key findings/state of knowledge Nitrogen cycling and nitrogen stocks in terrestrial ecosystems significantly differ between different ecosystem types (arable, grassland, shrubland, forests). Nitrogen stocks of managed systems are increased by fertilization and N retention processes are negatively affected. It is also obvious that nitrogen processes in natural and semi-natural ecosystems have already been affected by atmospheric N r input. Following perturbations of the N cycle, terrestrial ecosystems are increasingly losing N via nitrate leaching and gaseous losses (N 2 O, NO, N 2 and in agricultural systems also NH 3 ) to the environment.

Nitrogen mineralization and denitrification as influenced by crop residue particle size

Managing the crop residue particle size has the potential to affect N conservation in agricultural systems. We investigated the influence of barley (Hordeum vulgare) and pea (Pisum sativum) crop residue particle size on N mineralization and denitrification in two laboratory experiments. Experiment 1: 15N-labelled ground (≤3 mm) and cut (25 mm) barley residue, and microcrystalline cellulose+glucose were mixed into a sandy loam soil with additional inorganic N. Experiment 2: inorganic15 N and C2H2 were added to soils with barley and pea material after 3, 26, and 109 days for measuring gross N mineralization and denitrification.Net N immobilization over 60 days in Experiment 1 cumulated to 63 mg N kg-1 soil (ground barley), 42 (cut barley), and 122 (cellulose+glucose). More N was seemingly net mineralized from ground barley (3.3 mg N kg-1 soil) than from cut barley (2.7 mg N kg-1 soil). Microbial biomass peaked at day 4 with the barley treatments and at day 14 with the cellulose+glucose whereafter the biomass leveled out at values 79 mg C kg-1 (ground), 104 (cut), and 242 (cellulose+glucose) higher than for the control soil. Microbial growth yields were similar for the two barley treatments, ca. 60 mg C g-1 substrate C added, which was lower than the 142 mg C g-1 C added with cellulose+glucose. This suggests that the 75% (w/w) holocelluloses and sugars contained with the barley material remained physically protected despite grinding. In Experiment 2 gross mineralization on day 3 was 4.8 mg N kg-1 d-1 with ground pea, twice as much as for all other treatments. On day 26 the treatment with ground barley had the greatest gross N mineralization. In static cores ground barley denitrified 11-fold more than did cut barley, whereas denitrification was similar for the two pea treatments. In suspensions denitrification was similar for the two treatments both with barley and pea residue.We conclude that the higher microbial activity associated with the initial decomposition of ground plant material is due to a more intimate plant residue-soil contact. On the long term, grinding the plant residues has no significant effect on N dynamics.

Nitrous oxide emission from an agricultural field : comparison between measurements by flux chamber and micrometerological techniques

The soil in a drained fjord area, reclaimed for arable farming, produced N2O mainly at 75–105 cm depth, just above the ground water level. Surface emissions of N2O were measured from discrete small areas by closed and open-flow chamber methods, using gas chromatographic analysis and over larger areas by integrative methods: flux gradient (analysis by FTIR), conditional sampling (analysis by TDLAS), and eddy covariance (analysis by TDLAS). The mean emission of N2O as determined by chamber procedures during a 9-day campaign was 162–202 μg N2ONm−2h−1 from a wheat stubble and 328–467 μg N2ONm−2 h−1 from a carrot field. The integrative approaches gave N2O emissions of 149–495 μg N2ONm−2 h−1, i.e. a range similar to those determined with the chamber methods. Wind direction affected the comparison of chamber and integrative methods because of patchiness of the N2O emission over the area. When a uniform area with a single type of vegetation had a dominant effect on the N2O gradient at the sampling mast, the temporal variation in N2O emission determined by the flux gradient/FTIR method and chamber methods was very similar, with differences of only 18% or less in mean N2O emission, well below the variation encountered with the chamber methods themselves. A detailed comparison of FTIR gradient and chamber data taking into account the precise emission footprint showed good agreement. It is concluded that there was no bias between the different approaches used to measure the N2O emission and that the precision of the measurements was determined by the spatial variability of the N2O emission at the site and the variability inherent in the individual techniques. These results confirm that measurements of N2O emissions from different ecosystems obtained by the different methods can be meaningfully compared.

Denitrification variability and control in a riparian fen irrigated with agricultural drainage water

Abstract Denitrification was measured by the C 2 H 2 inhibition technique in a riparian fen irrigated with agricultural drainage water. 16 h after C 2 H 2 treatment 88 ± 14% of the total N 2 O contained in water-saturated cores could be accounted for by assuming equilibrium between the gas phase and the liquid phase. The denitrification activity averaged 2.8 and 8.8 mg N 2 O-N m −2 day −1 in the control plot and 1.6 and 21.9mg N 2 O-N m −2 day −1 in the irrigated plot during the dry and the runoff periods respectively. Four percent of the incoming NO 3 − was reduced to gaseous N. The spatial variability was often high, with coefficients of variation > 100% and was independent of seasonal changes in soil anaerobiosis. Soil NO 3 − and denitrification were poorly related, and bulk concentrations of NO 3 − below 200 μm suggested that the process was strongly limited by diffusion of NO 3 − into the soil during periods of flooding. Mean denitrification and water-filled pores correlated positively, r = 0.71∗∗∗ for the control and r = 0.68∗∗∗ for the irrigated plots. Water-soluble C was not related to denitrification. Multiple regression models including soil water, NO 3 − , soluble C and temperature as independent variables, predicted between 21 and 55% of the denitrification, the highest value found when only mean data was considered. Water-filled pores was the most important variable. The observations on which 2 variables controlled denitrification were supported by laboratory experiments with manipulated cores. Water additions increased denitrification only in samples collected during the dry period. Anaerobic incubation of saturated cores did not affect the process. Restricted NO 3 − availability was clearly illustrated by the 25–41-fold increase obtained when NO 3 − was injected into cores at ambient and high carbon respectively. A response of up to 13-fold was observed when substrate-amended cores were made into slurries. Glucose did not increase denitrification by more than a factor of three.

Terrestrial plant methane production and emission

In this minireview, we evaluate all experimental work published on the phenomenon of aerobic methane (CH(4) ) generation in terrestrial plants and plant. Clearly, despite much uncertainty and skepticism, we conclude that the phenomenon is true. Four stimulating factors have been observed to induce aerobic plant CH(4) production, i.e. cutting injuries, increasing temperature, ultraviolet radiation and reactive oxygen species. Further, we analyze rates of measured emission of aerobically produced CH(4) in pectin and in plant tissues from different studies and argue that pectin is very far from the sole contributing precursor. In consequence, scaling up of aerobic CH(4) emission needs to take into consideration other potential sources than pectin. Due to the large uncertainties related to effects of stimulating factors, genotypic responses and type of precursors, we conclude that current attempts for upscaling aerobic CH(4) into a global budget is inadequate. Thus it is too early to draw the line under the aerobic methane emission in plants. Future work is needed for establishing the relative contribution of several proven potential CH(4) precursors in plant material.