Models have been formulated for monospecific stands in which canopy photosynthesis is determined by the
vertical distribution of leaf area, nitrogen and light. In such stands, resident plants can maximize canopy
photosynthesis by distributing their nitrogen parallel to the light gradient, with high contents per unit leaf area
at the top of the vegetation and low contents at the bottom. Using principles from game theory, we expanded these
models by introducing a second species into the vegetation, with the same vertical distribution of biomass and
nitrogen as the resident plants but with the ability to adjust its specific leaf area (SLA, leaf area[ratio ]leaf mass). The
rule of the game is that invaders replace the resident plants if they have a higher plant carbon gain than those of
the resident plants. We showed that such invaders induce major changes in the vegetation. By increasing their
SLA, invading plants could increase their light interception as well as their photosynthetic nitrogen-use efficiency
(PNUE, the rate of photosynthesis per unit organic nitrogen). By comparison with stands in which canopy
photosynthesis is maximized, those invaded by species of high SLA have the following characteristics: (1) the leaf
area index is higher; (2) the vertical distribution of nitrogen is skewed less; (3) as a result of the supra-optimal
leaf area index and the more uniform distribution of nitrogen, total canopy photosynthesis is lower. Thus, in
dense canopies we face a classical tragedy of the commons: plants that have a strategy to maximize canopy carbon
gain cannot compete with those that maximize their own carbon gain. However, because of this strategy, individual
as well as total canopy carbon gain are eventually lower. We showed that it is an evolutionarily stable strategy
to increase SLA up to the point where the PNUE of each leaf is maximized.