Trifolium repens L. and Lolium perenne L. were grown in monocultures and bi‐species mixture in a Free Air Carbon Dioxide Enrichment (FACE) experiment at elevated (60 Pa) and ambient (35 Pa) CO2 partial pressure (pCO2) for three years. The effects of defoliation frequencies (4 and 7 cuts in 1993; 4 and 8 cuts in 1994/95) and nitrogen fertilization (10 and 42 g m–2 y–1 N in 1993; 14 and 56 g m–2 y–1 N in 1994/95) on the growth response to pCO2 were investigated. There were significant interspecific differences in the CO2 responses during the first two years, while in the third growing season, these interspecific differences disappeared. Yield of T. repens in monocultures increased in the first two years by 20% when grown at elevated pCO2. This CO2 response was independent of defoliation frequency and nitrogen fertilization. In the third year, the CO2 response of T. repens declined to 11%. In contrast, yield of L. perenne monocultures increased by only 7% on average over three years at elevated pCO2. The yield response of L. perenne to CO2 changed according to defoliation frequency and nitrogen fertilization, mainly in the second and third year. The ratio of root/yield of L. perenne increased under elevated pCO2, low N fertilizer rate, and frequent defoliation, but it remained unchanged in T. repens. We suggest that the more abundant root growth of L. perenne was related to increased N limitation under elevated pCO2. The consequence of these interspecific differences in the CO2 response was a higher proportion of T. repens in the mixed swards at elevated pCO2. This was evident in all combinations of defoliation and nitrogen treatments. However, the proportion of the species was more strongly affected by N fertilization and defoliation frequency than by elevated pCO2. Based on these results, we conclude that the species proportion in managed grassland may change as the CO2 concentration increases. However, an adapted management could, at least partially, counteract such CO2 induced changes in the proportion of the species. Since the availability of mineral N in the soil may be important for the species’ responses to elevated pCO2, more long‐term studies, particularly of processes in the soil, are required to predict the entire ecosystem response.

~Symbiotic N, fixation is one of the main processes that introduces N into terrestrial ecosystems. As such, it may be crucial for the sequestration of the extra C available in a world of continuously increasing atmospheric CO, partial pressure (pC0,). The effect of elevated pC0, (60 Pa) on symbiotic N, fixation ('5N-isotope dilution method) was investigated using Free-Air-C0,-Enrichment technology over a period of 3 years. Trifolium repens was cultivated either alone or together with Lolium perenne (a nonfixing reference crop) in mixed swards. Two different N fertilization levels and defoliation frequencies were applied. The total N yield increased consistently and the percentage of plant N derived from symbiotic N, fixation increased significantly in T. repens under elevated pC0,. All additionally assimilated N was derived from symbiotic N, fixation, not from the soil. In the mixtures exposed to elevated pCO,, an increased amount of symbiotically fixed N (+7.8,8.2, and 6.2 g m-' a-' in 1993, 1994, and 1995, respectively) was introduced into the system. lncreased N, fixation is a competitive advantage for T. repens in mixed swards with pasture grasses and may be a crucial factor in maintaining the C:N ratio in the ecosystem as a whole.

The extent of the response of plant growth to atmospheric CO enrichment depends on the availability of resources other than CO. An important growth-limiting resource under field conditions is nitrogen (N). N may, therefore, influence the CO response of plants. The effect of elevated CO (60 Pa) partial pressure (pCO) on the N nutrition of field-grown Lolium perenne swards, cultivated alone or in association with Trifolium repens, was investigated using free air carbon dioxide enrichment (FACE) technology over 3 years. The established grassland ecosystems were treated with two N fertilization levels and were defoliated at two frequencies. Under elevated pCO, the above-ground plant material of the L. perenne monoculture showed a consistent and significant decline in N concentration which, in general, led to a lower total annual N yield. Despite the decline in the critical N concentration (minimum N concentration required for non-N-limited biomass production) under elevated pCO, the index of N nutrition (ratio of actual N concentration and critical N concentration) was lower under elevated pCO than under ambient pCO in frequently defoliated L. perenne monocultures. Thus, we suggest that reduced N yield under elevated pCO was evoked indirectly by a reduction of plant-available N. For L. perenne grown in association with T. repens and exposed to elevated pCO, there was an increase in the contribution of symbiotically fixed N to the total N yield of the grass. This can be explained by an increased apparent transfer of N from the associated N-fixing legume species to the non-fixing grass. The total annual N yield of the mixed grass/legume swards increased under elevated pCO. All the additional N yielded was due to symbiotically fixed N. Through the presence of an N-fixing plant species more symbiotically fixed N was introduced into the system and consequently helped to overcome N limitation under elevated pCO.

The effect of an elevated partial pressure of CO2 (pCO2) on carbohydrate concentrations in source leaves and pseudo‐stems (stubble) of Lolium perenne L. (perennial ryegrass) during regrowth was studied in a regularly defoliated grass sward in the field. The free air carbon dioxide enrichment (FACE) technology enabled natural environmental conditions to be provided. Two levels of nitrogen (N) supply were used to modulate potential plant growth. Carbohydrate concentrations in source leaves were increased at elevated pCO2, particularly at low N supply. Elevated leaf carbohydrate concentrations were related to an increased structural carbon (C) to N ratio and thus reflected an increased C availability together with a N‐dependent sink limitation. Immediately after defoliation, apparent assimilate export rates (differences in the carbohydrate concentrations of young source leaves measured in the evening and on the following morning) showed a greater increase at elevated pCO2 than at ambient pCO2; however, replenishment of carbohydrate reserves was not accelerated. Distinct, treatment‐dependent carbohydrate concentrations in pseudo‐stems suggested an increasing degree of C‐sink limitation from the treatment at ambient pCO2 with high N supply to that at elevated pCO2 with low N supply. During two growing seasons, no evidence of a substantial change in the response of the carbohydrate source in L. perenne to elevated pCO2 was found. Our results support the view that the response of L. perenne to elevated pCO2 is restricted by a C‐sink limitation, which is particularly severe at low N supply.

Lolium perenne and Trifolium repens were grown in a Free Air CO2 Enrichment (FACE) system at elevated (600 }.imol mol"*) and ambient (340 [jmol mol"^) carbon dioxide concentrations during a whole growing season. Using a root ingrowth bag technique the extent to which CO2 enrichment influenced the growth of L, perenne and T. repens roots under two contrasting nutrient regimes was examined. Root ingrowth bags were inserted for a fixed time into the soil in order to trap roots. It was also possible to follow the mortality of roots in bags inserted for different time intervals. Rool ingrowth of both L. perenne and T. repens increased under elevated COj conditions. In L. perenne, root ingrowth decreased with increasing nutrient fertilizer level, but for T. repens the root ingrowth was not affected by the nutrient application rate. Besides biomass measurements, root length estimates were made for T, repens. These showed an increase under elevated CO2 concentrations. Root decomposition appeared to decrease under elevated CO2 concentrations. A possible explanation for this effect is the observed changes in tissue composition, such as the increase in the carbonmitrogen ratio in roots of L. perenne at elevated CO2 concentrations.

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