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Elevated CO₂ concentrations were found to cause early senescence during leaf development in sunflower (Helianthus annuus L.) plants, probably by reducing nitrogen availability since key enzymes of nitrogen metabolism, including nitrate reductase (NR); glutamine synthetase (GS) and glutamate dehydrogenase (GDH), were affected. Elevated CO₂ concentrations significantly decreased the activity of nitrogen assimilation enzymes (NR and GS) and increased GDH deaminating activities. Moreover, they substantially rose the transcript levels of GS1 while lowering those of GS2. Increased atmospheric CO₂ concentrations doubled the CO₂ fixation and increased transpiration rates, although these parameters decreased during leaf ontogeny. It can be concluded that elevated atmospheric CO₂ concentrations alter enzymes involved in nitrogen metabolism at the transcriptional and post-transcriptional levels, thereby boosting mobilization of nitrogen in leaves and triggering early senescence in sunflower plants.

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Different parameters that vary during leaf development may be affected by light intensity. To study the influence of different light intensities on primary leaf senescence, sunflower (Helianthus annuus L.) plants were grown for 50 days under two photon flux density (PFD) conditions, namely high irradiance (HI) at 350 mol(photon) m -2 s -1 and low irradiance (LI) at 125 mol(photon) m -2 s -1 . Plants grown under HI exhibited greater specific leaf mass referred to dry mass, leaf area and soluble protein at the beginning of the leaf development. This might have resulted from the increased CO 2 fixation rate observed in HI plants, during early development of primary leaves. Chlorophyll a and b contents in HI plants were lower than in LI plants in young leaves. By contrast, the carotenoid content was significantly higher in HI plants. Glucose concentration increased with the leaf age in both treatments (HI and LI), while the starch content decreased sharply in HI plants, but only slightly in LI plants. Glucose contents were higher in HI plants than in LI plants; the differences were statistically significant (p<0.05) mainly at the beginning of the leaf senescence. On the other hand, starch contents were higher in HI plants than in LI plants, throughout the whole leaf development period. Nitrate reductase (NR) activity decreased with leaf ageing in both treatments. However, the NR activation state was higher during early leaf development and decreased more markedly in senescent leaves in plants grown under HI. GS activity also decreased during sunflower leaf ageing under both PFD conditions, but HI plants showed higher GS activities than LI plants. Aminating and deaminating activities of glutamate dehydrogenase (GDH) peaked at 50 days (senescent leaves). GDH deaminating activity increased 5-fold during the leaf development in HI plants, but only 2-fold in LI plants. The plants grown under HI exhibited considerable oxidative stress in vivo during the leaf senescence, as revealed by the substantial H 2 O 2 accumulation and the sharply decrease in the antioxidant enzymes, catalase and ascorbate peroxidase, in comparison with LI plants. Probably, systemic signals triggered by a high PFD caused early senescence and diminished oxidative protection in primary leaves of sunflower plants as a result.