Concepts of the regulation of assimilate partitioning in leaves frequently consider only the allocation of carbon between sucrose and starch synthesis, storage and export. While carbohydrate metabolism accounts for a large proportion of assimilated carbon, such analyses provide only a restricted view of carbon metabolism and partitioning in leaf cells since photosynthetic carbon fixation provides precursors for all other biosynthetic pathways in the plant. Most of these precursors are required for biosynthesis of amino acids that form the building blocks for many compounds in plants. We have used leaf carbon : nitrogen ratios to calculate the allocation of photosynthetic electrons to the assimilation of nitrogen necessary for amino acid formation, and conclude that this allocation is variable but may be higher than values often quoted in the literature. Respiration is a significant fate of fixed carbon. In addition to supplying biosynthetic precursors, respiration is required for energy production and may also act, in both light and dark, to balance cellular energy budgets. We have used growth CO2 concentration and irradiance to modify source activity in Lolium temulentum in order to explore the interactions between photosynthetic carbon and nitrogen assimilation, assimilate production, respiration and export. It is demonstrated that there is a robust correlation between source activity and foliar respiration rates. Under some conditions concomitant increases in source activity and respiration may be necessary to support faster growth. In other conditions, increases in respiration appear to result from internal homeostatic mechanisms that may be candidate targets for increasing yield.

Lolium temulentum L. Ba 3081 was grown hydroponically in air (350 &mgr;mol mol(-1) CO(2)) and elevated CO(2) (700 &mgr;mol mol(-1) CO(2)) at two irradiances (150 and 500 &mgr;mol m(-2) s(-1)) for 35 days at which point the plants were harvested. Elevated CO(2) did not modify relative growth rate or biomass at either irradiance. Foliar carbon-to-nitrogen ratios were decreased at elevated CO(2) and plants had a greater number of shorter tillers, particularly at the lower growth irradiance. Both light-limited and light-saturated rates of photosynthesis were stimulated. The amount of ribulose-1, 5-bisphosphate carboxylase-oxygenase (Rubisco) protein was increased at elevated CO(2), but maximum extractable Rubisco activities were not significantly increased. A pronounced decrease in the Rubisco activation state was found with CO(2) enrichment, particularly at the higher growth irradiance. Elevated-CO(2)-induced changes in leaf carbohydrate composition were small in comparison to those caused by changes in irradiance. No CO(2)-dependent effects on fructan biosynthesis were observed. Leaf respiration rates were increased by 68% in plants grown with CO(2) enrichment and low light. We conclude that high CO(2) will only result in increased biomass if total light input favourably increases the photosynthesis-to-respiration ratio. At low irradiances, biomass is more limited by increased rates of respiration than by CO(2)-induced enhancement of photosynthesis.

This research was performed under the auspices of the U.S. Department of Energy under Contract No. DE-AC02-76CH00016.

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