Summary After a step increase in the atmospheric partial pressure of CO2 (pCO2), the availability of mineral N may be insufficient to meet the plant's increased demand for N. Over time, however, the ecosystem may adapt to the new conditions, and a new equilibrium may be established in the fluxes of C and N. This would result in a higher dry mass (DM) yield response of the plants to elevated pCO2. The effect of elevated atmospheric pCO2 (60 Pa pCO2) was studied in Lolium perenne L. swards with two N fertilization treatments (14 and 56 g m−2 y−1) in a six‐year FACE (Free Air Carbon dioxide Enrichment) experiment. In the high N treatment, the input of N with fertilizer considerably exceeded the export of N with the harvested plant material in both CO2 treatments leading to an apparent net input of N into the ecosystem. Accordingly, the proportion of harvested N derived from 15N labelled fertilizer N, applied throughout the experiment (< 6 years), increased over the years. Under these high N conditions, the annual DM yield response of the Lolium perenne sward to elevated pCO2 increased (from 7% in 1993 to 25% in 1998). In parallel, the response of N yield to elevated pCO2 increased, and the initially negative effect of elevated pCO2 on specific leaf area (SLA) disappeared. The high N input system seemed to overcome in part an initially limiting effect of N on the yield response to elevated pCO2 within a few years. In contrast, there was no apparent net input of N into the ecosystem in the low N treatment, because N fertilization just compensated the export of N with the harvested plant material. Accordingly, the proportion of harvested N yield, derived from fertilizer N, which was applied throughout the experiment, remained low. At low N, the availability of mineral N strongly limited plant growth and yield production in both CO2 treatments; the low yields of DM and N, the low concentration of N in the plant material, and the low SLA reflected this. Although the plants grew under the same environmental conditions and the same management treatment as plants in the high N treatment, the response of DM yields to elevated pCO2 in the low N treatment remained weak throughout the experiment (5% in 1993 and 9% in 1998). The results are discussed in the context of the sizes of the different N pools in the soil, the allocation of N within the plant and the possible effects on temporal immobilization, and the availability of mineral N for yield production as affected by elevated pCO2 and N fertilization.
Lolium perenne, a main component species in managed grassland, is well adapted to defoliation, fertilization, and regrowth cycles; and hence, to changes in the assimilatory carbon source‐sink ratio. In the Swiss Free Air CO2 Enrichment experiment the source‐sink ratio is (i) increased by elevated partial pressure of CO2 (pCO2), (ii) decreased by enhanced carbon use under high N fertilization, and (iii) gradually increased during regrowth after defoliation. Since sucrose synthesis plays a central role in leaf carbohydrate metabolism in this fructan‐accumulating species, we investigated how sucrose‐phosphate synthase (SPS) responds to the differing assimilatory carbon fluxes and source‐sink ratios in the field. Assimilatory carbon flux, as estimated by leaf gas exchange, strongly depended on pCO2. Surprisingly, the SPS content per leaf area did not increase with pCO2, but increased with N fertilization. During later regrowth, when a dense canopy had formed, the SPS content decreased; in particular, SPS was decreased at high N under elevated pCO2. Further, the higher assimilatory carbon flux through SPS at elevated pCO2 was accompanied by a higher activation state of SPS. The SPS content correlated very strongly with the ratio of free sucrose to free amino acid in leaves, which represents the carbon source‐sink ratio. Hence, SPS content in L. perenne appears to be regulated by the current, strongly nitrogen‐dependent, source‐sink relation.
This paper originates from a presentation at the International Conference on Assimilate Transport and Partitioning, Newcastle, NSW, August 1999 Swards of Lolium perenne L. were grown in the field in a long-term free air CO2 enrichment (FACE) facility. The CO2 treatment was combined with two levels of N fertilization and regular defoliation, which resulted in plants with a wide range of source–sink relations. C and N metabolism were investigated to assess the role of carbohydrate and nitrogenous compounds in leaves in indicating source–sink relations. Sucrose exhibited the largest changes in contents during the day–night cycle; therefore, it was identified as the main short-term storage compound for night-time export. Fructan accumulation indicated the degree of surplus C supply in the source compared to C use in sinks. Nitrate content depended mainly on N fertilization, and was reduced under elevated pCO2. Nitrate appeared to indicate a current surplus of available N relative to the need for growth. Amino acid content responded strongly to N fertilization but decreased only slightly under elevated pCO2. Protein content, however, decreased significantly under elevated pCO2. The patterns of diurnal changes of C or N compounds did not differ between CO2 treatments. Down-regulation of photosynthesis appeared to occur when plants were extremely N-limited as under elevated pCO2, low N and at a late regrowth stage.
Abstract.To investigate the importance of phosphorus and carbohydrate concentrations in influencing photosynthetic capacity of tropical forest tree seedlings under elevated CO 2 , we grew seedlings of Beilschmiedia pendula (Sw.) Hemsl. (Lauraceae) under elevated CO 2 concentrations either with or without vesicular-arbuscular (VA) mycorrhizae. VA-mycorrhizae increased phosphorus concentrations in all plant organs (leaves, stems and roots). Maximum rates of photosynthesis (A max ) measured under saturating levels of CO 2 and light were correlated with leaf phosphorus concentrations. VA-mycorrhizae also increased leaf carbohydrate concentrations, particularly under elevated CO 2 , but levels were low and within the range observed in naturally occurring forest species. Root carbohydrate concentrations were reduced in VA-mycorrhizal plants relative to non-mycorrhizal plants. These results indicate an important role for VA-mycorrhizae in controlling photosynthetic rates and sink strength in tropical trees, and thus in determining their response to future increases in atmospheric CO 2 concentrations.
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