Effects of nitrogen supply on the anatomy and chemical composition of leaves of four grass species belonging to the genus Poa, as determined by image-processing analysis and pyrolysis–mass spectrometry


H. Lambers, Department of Plain Ecology and Evolutionary Biology, University of Utrecht, P.O. Box 800-84 3508 TB Utrecht, The Netherlands.


Previous experiments have shown that the anatomy and chemical composition of leaves of inherently fast- and slow-growing grass species, grown at non-limiting nitrogen supply, differ systematically. The present experiment was carried out to investigate whether these differences persist when the plants are grown at an intermediate or a very low nitrogen supply. To this end, the inherently fast-growing Poa annua L. and Poa trivialis L., and the inherently slow-growing Poa compressa L. and Poa pratensis (L.) Schreb. were grown hydroponically at three levels of nitrate supply: at optimum (RGRmax) and at relative addition rates of 100 and 50 mmol N (mol N)−1 d−1 (RAR100 and RAR50), respectively.

As expected, at the lowest N supply, the potentially fast-growing species grew at the same rate as the inherently slow-growing ones. Similarly, the differences in leaf area ratio (LAR, leaf area:total dry mass), specific leaf area (SLA, leaf arear:leaf dry mass) and leaf mass ratio (LMR, leaf dry mass:total dry mass) disappeared. Under optimal conditions, the fast-growing species differed from the slow-growing ones in that they had a higher N concentration. There were no significant differences in C concentration. With decreasing N supply, the total N concentration decreased and the differences between the species disappeared. The total C concentration increased for the fast-growing species and decreased for the slow-growing ones, i.e. the small, but insignificant, difference in C concentration between the species at RGRmax increased with decreasing N supply.

The chemical composition of the leaves at low N supply, analysed in more detail by pyrolysis–mass spectrometry, showed an increase in the relative amounts of guaiacyl lignin, cellulose and hemicellulose, whereas those of syringyl lignin and protein decreased.

The anatomy and morphology of the leaves of the four grass species differing in RGRmax were analysed by image-processing analysis. The proportion of the total volume occupied by mesophyll plus intercellular spaces and epidermis did not correlate with the amount of leaf mass per unit leaf area (specific leaf mass, SLM) at different N supply. The higher SLM at low N supply was caused partly by a high proportion of non-veinal sclerenchymatic cells per cross-section and partly by the smaller volume of epidermal cells.

We conclude that the decrease in relative growth rate (and increase in SLM) at decreasing N supply is partly due to chemical and anatomical changes. The differences between the fast- and slow-growing grass species at an optimum nutrient supply diminished when plants were growing at a limiting nitrogen supply.