Plant Responses to Multiple Environmental FactorsPhysiological ecology provides tools for studying how interacting environmental resources control plant growth

M ost plants require a similar balance of resources-energy, water, and mineral nutrients-to maintain optimal growth. Natural environments, however, differ by at least two orders of magnitude in the availability of these resources. Light intensity varies 100fold from the canopy to the floor of a rainforest (Bj6rkman 1981); annual precipitation ranges 500-fold (105000 mm/yr) from deserts to tropical rainforests; and the amount of nitrogen available to plants varies from 0.09 g/m2 * yr in polar desert (Dowding et al. 1981) to 22.8 g/m2 * yr in a rich tropical rainforest (Vitousek 1984). Plants growing in these diverse environments maintain tissue concen-

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