A model of daily canopy photosynthesis was constructed taking light and leaf nitrogen distribution in the canopy into consideration. It was applied to a canopy of Solidago altissima. Both irradiance and nitrogen concentration per unit leaf area decreased exponentially with increasing cumulative leaf area from the top of the canopy. The photosynthetic capacity of a single leaf was evaluated in relation to irradiance and nitrogen concentration. By integration, daily canopy photosynthesis was calculated for various canopy architectures and nitrogen allocation patterns. The optimal pattern of nitrogen distribution that maximizes the canopy photosynthesis was determined. Actual distribution of leaf nitrogen in the canopy was more uniform than the optimal one, but it realized over 20% more photosynthesis than that under uniform distribution and 4.7% less photosynthesis than that under the optimal distribution. Redeployment of leaf nitrogen to the top of the canopy with ageing should be more effective in increasing total canopy photosynthesis in a stand with a dense canopy than in a stand with an open canopy.
Photosynthetic nitrogen use efficiency (PNUE, photosynthetic capacity per unit leaf nitrogen) is one of the most important factors for the interspecific variation in photosynthetic capacity. PNUE was analysed in two evergreen and two deciduous species of the genus Quercus . PNUE was lower in evergreen than in deciduous species, which was primarily ascribed to a smaller fraction of nitrogen allocated to the photosynthetic apparatus in evergreen species. Leaf nitrogen was further analysed into proteins in the water-soluble, the detergent-soluble, and the detergentinsoluble fractions. It was assumed that the detergentinsoluble protein represented the cell wall proteins. The fraction of nitrogen allocated to the detergent-insoluble protein was greater in evergreen than in deciduous leaves. Thus the smaller allocation of nitrogen to the photosynthetic apparatus in evergreen species was associated with the greater allocation to cell walls. Across species, the fraction of nitrogen in detergent-insoluble proteins was positively correlated with leaf mass per area, whereas that in the photosynthetic proteins was negatively correlated. There may be a trade-off in nitrogen partitioning between components pertaining to productivity (photosynthetic proteins) and those pertaining to persistence (structural proteins). This trade-off may result in the convergence of leaf traits, where species with a longer leaf life-span have a greater leaf mass per area, lower photosynthetic capacity, and lower PNUE regardless of life form, phyllogeny, and biome.
There is a strong correlation between leaf thickness and the light-saturated rate of photosynthesis per unit leaf area ( P max ). However, when leaves are exposed to higher light intensities after maturation, P max often increases without increasing leaf thickness. To elucidate the mechanism with which mature leaves increase P max , the change in anatomical and physiological characteristics of mature leaves of Chenopodium album, which was transferred from low to high light condition, were examined. When compared with leaves subjected to low light continuously (LL leaves), the leaves transferred from low to high light (LH leaves) significantly increased P max . The transfer also increased the area of chloroplasts facing the intercellular space ( S c ) and maintained a strong correlation between P max and S c . The mesophyll cells of LL leaves had open spaces along cell walls where chloroplasts were absent, which enabled the leaves to increase P max when they were exposed to high light (LH). However, the LH leaves were not thick enough to allow further increase in P max to the level in HH leaves. Thus leaf thickness determines an upper limit of P max of leaves subjected to a change from low to high light conditions. Shade leaves would only increase P max when they have open space to accommodate chloroplasts which elongate after light conditions improve.
Summary1. Nitrogen (N) is an essential limiting resource for plant growth, and its efficient use may increase fitness. We investigated photosynthetic N-use efficiency (photosynthetic capacity per unit N) in relation to N allocation to Rubisco and to cell walls in Polygonum cuspidatum Sieb. et Zucc. which germinated in May (early germinators) and August (late germinators). 2. There was a significant difference between early and late germinators in photosynthetic capacity as a function of leaf N content per unit area. Higher photosynthetic N-use efficiency in late germinators was caused primarily by a larger allocation of N to Rubisco. 3. Nitrogen allocation to cell walls was smaller in late germinators. The shorter growth period in late germinators was associated with higher photosynthetic capacity, which was achieved by allocating more N to photosynthetic proteins at the expense of cell walls. 4. The trade-off between N allocation to photosynthesis and to structural tissues suggests that plants change N allocation to increase either the rate or duration of carbon assimilation. Such plastic change would help plants maintain themselves and cope with environmental changes.
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