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The effect of elevated (60 Pa) atmospheric carbon dioxide partial pressure (pCO2) and N fertilisation on the availability of mineral N and on N transformation in the soil of a Lolium perenne L. monoculture was investigated in the Swiss FACE (Free Air Carbon dioxide Enrichment) experiment. The apparent availability of nitrate and ammonium for plants was estimated during a representative, vegetative re-growth period at weekly intervals from the sorption of the minerals to mixed-bed ion-exchange resin bags at a soil depth of 5 cm. N mineralisation was measured using sequential coring and in situ exposure of soil cores in the top 10 cm of the soil before and after the first cut in spring 1997. High amounts of mineral N were bound to the ion exchange resin during the first week of re-growth. This was probably the combined result of the fertiliser application, the weak demand for N by the newly cut sward and presumably high rates of root decay and exudation after cutting the sward. During the first 2 weeks after the application of fertiliser N at the first cut, there was a dramatic reduction in available N; N remained low during the subsequent weeks of re-growth in all treatments. Overall, nitrate was the predominant form of mineral N that bound to the resin for the duration of the experiment. Apparently, there was always more nitrate than ammonium available to the plants in the high N fertilisation treatment for the whole re-growth period. Apparent N availability was affected significantly by elevated pCO2 only in the first week after the cut; under high N fertilisation, elevated pCO2 increased the amount of mineral N that was apparently available to the plants. Elevated pCO2 did not affect apparent net transformation of N, loss of N or uptake of N by plants. The present data are consistent with earlier results and suggest that the amount of N available to plants from soil resources does not generally increase under elevated atmospheric pCO2. Thus, a possible limiting effect of N on primary production could become more stringent under elevated atmospheric pCO2 as the demand of the plant for N increases.

Experimental findings indicate that, in terrestrial ecosystems, nitrogen cycling changes under elevated partial pressure of atmospheric CO2 (pCO2). It was suggested that the concentration of N in plant litter as well as the amount of litter are responsible for these changes. However, for grassland ecosystems, there have been no relevant data available to support this hypothesis. Data from five years of the Swiss FACE experiment show that, under fertile soil conditions in a binary plant community consisting of Lolium perenne L. and Trifolium repens L., the concentration of litter N does not change under elevated atmospheric pCO2; this applies to harvest losses, stubble, stolons and roots as the sources of litter. This is in strong contrast to the CO2 response of L. perenne swards without associated legumes; in this case the above-ground concentration of biomass N decreased substantially. Increased symbiotic N2 fixation in T. repens nodules and a greater proportion of the N-rich T. repens in the community are regarded as the main mechanisms that buffer the increased C introduction into the ecosystem under elevated atmospheric pCO2. Our data also suggest that elevated atmospheric pCO2 results in greater amounts of litter, mainly due to increased root biomass production. This study indicates that, in a fertile grassland ecosystem with legumes, the concentration of N in plant litter is not affected by elevated atmospheric pCO2 and, thus, cannot explain CO2-induced changes in the cycling of N.

The significance of symbiotic N2 fixation in legumes (Trifolium alpinum L., T. nivale Sieber, T. pratense L., T. badium Schreber, T. thalii Vill., T. repens L., Lotus alpinus [DC.] Schleicher, L. corniculatus L., Vicia sativa L.) and other N sources for the N budget of grassland ecosystems was studied along an altitudinal gradient in the Swiss Alps. The total annual symbiotic N2 fixation was compared with other sources of N for plant growth of the total plant community (mineralisation and wet deposition). The contribution of symbiotically fixed N to total above-ground N yield of the swards decreased from at least 16% to 9% with increasing altitude where legumes were present. This decrease was due to a decrease in the yield proportion of legumes from 15% at 900 and 1380 m a.s.l. to 5% at 2100 and 2300 m a.s.l. (no legumes were found above 2750 m a.s.l.) and not to a decline in the activity of symbiotic N2 fixation. With increasing altitude legumes are more patchily distributed. The high symbiotic N2 fixation of individual plants up to their altitudinal limit is not primarily the result of low mineral N availability since an addition of NH4 + or NO3 − fertiliser at 2300 m a.s.l. led either to no decrease or only to a minor decrease in symbiotic N2 fixation. At 1380 m a.s.l., N mineralisation (13.45 g N m−2 yr−1) appeared to be the main source of N for growth of the sward; N from symbiosis (at least 1.0 g to 2.6 g N m−2 yr−1) and wet deposition (0.4 g to 0.6 g m−2 yr−1) was not a significant N source for plant growth at this altitude. At 2100 m a.s.l., the combined amounts of N from symbiotic N2 fixation (at least 0.1 g N m−2 yr−1) and wet deposition (0.3 g N m−2 yr−1) appeared to be similarly important for plant growth as soil N mineralisation (0.47 g N m−2 yr−1). At high altitudes, wet N deposition and symbiotic N2 fixation together represent a significant source of N for the grassland ecosystem while at low altitudes these N inputs appear to be much less important.

The growth response of white clover (Trifolium repens L.) to the expected increase in atmospheric partial pressure of CO2 (pCO2) may depend on P availability. A decrease in the rate of transpiration due to increased pCO2 may reduce the amount of P transported to the shoot, thereby causing a change in the partitioning of P between the root and shoot. To test these hypotheses, four concentrations of P in the nutrient solution, combined with two pCO2 treatments, were applied to nodulated white clover plants. Compared to ambient pCO2 (35 Pa), twice ambient pCO2 (70 Pa) reduced the rate of transpiration but did not impair the total P uptake per plant. However, at twice ambient pCO2 and a moderate to high supply of P, concentrations of structural P and soluble P (Pi) were lower in the leaves and higher in the roots. The activity of root acid phosphatase was lower at twice ambient pCO2 than at ambient pCO2; it depended on the Pi concentration in the roots. At the highest P concentration, twice ambient pCO2 stimulated photosynthesis and the growth rate of the plant without affecting the concentration of nonstructural carbohydrates in the leaves. However, at the lower P concentrations, plants at twice ambient pCO2 lost their stimulation of photosynthesis in the afternoon, they accumulated nonstructural carbohydrates in the leaves and their growth rate was not stimulated; indicating C-sink limitation of growth. P nutrition will be crucial to the growth of white clover under the expected future conditions of increased pCO2.

Potassium (K) is reported to improve plant's resistance against environmental stress. A frequently experienced stress for plants in the tropics is water shortage. It is not known if sufficient K supply would help plants to partially overcome the effects of water stress, especially that of symbiotic nitrogen fixation which is often rather low in the tropics when compared to that of temperate regions. Thus, the impact of three levels of fertilizer potassium (0.1, 0.8 and 3.0 mM K) on symbiotic nitrogen fixation was evaluated with two legumes under high (field capacity to 25% depletion) and low (less than 50% of field capacity) water regimes. Plants were grown in single pots in silica sand under controlled conditions with 1.5 mM N (15N enriched NH4NO3). The species were faba bean (Vicia faba L.), a temperate, amide producing legume and common bean (Phaseolus vulgaris L.), a tropical, ureide producing species. In both species, 0.1 mM K was insufficient for nodulation at both moisture regimes, although plant growth was observed. The supply of 0.8 or 3.0 mM K allowed nodulation and subsequent nitrogen fixation which appeared to be adequate for respective plant growth. High potassium supply had a positive effect on nitrogen fixation, on shoot and root growth and on water potential in both water regimes. Where nodulation occurred, variations caused by either K or water supply had no consequences on the percentage of nitrogen derived from the symbiosis. The present data indicate that K can apparently alleviate water shortage to a certain extent. Moreover it is shown that the symbiotic system in both faba bean and common bean is less tolerant to limiting K supply than plants themselves. However, as long as nodulation occurs, N assimilation from the symbiotic source is not selectively affected by K as opposed to N assimilation from fertilizer.