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The ability to down-regulate leaf maximum net photosynthetic capacity (Amax) and dark respiration rate (Rdark) in response to shading is thought to be an important adaptation of trees to the wide range of light environments that they are exposed to across space and time. A simple, general rule that accurately described this down-regulation would improve carbon cycle models and enhance our understanding of how forest successional diversity is maintained. In this paper, we investigated the light response of Amax and Rdark for saplings of six temperate forest tree species in New Jersey, USA, and formulated a simple model of down-regulation that could be incorporated into carbon cycle models. We found that full-sun values of Amax and Rdark differed significantly among species, but the rate of down-regulation (proportional decrease in Amax or Rdark relative to the full-sun value) in response to shade was not significantly species- or taxon-specific. Shade leaves of sun-grown plants appear to follow the same pattern of down-regulation in response to shade as leaves of shade-grown plants. Given the light level above a leaf and one species-specific number (either the full-sun Amax or full-sun Rdark), we provide a formula that can accurately predict the leaf's Amax and Rdark. We further show that most of the down regulation of per unit area Rdark and Amax is caused by reductions in leaf mass per unit area (LMA): as light decreases, leaves get thinner, while per unit mass Amax and Rdark remain approximately constant.

The leaf economics spectrum (LES) describes multivariate correlations that constrain leaf traits of plant species primarily to a single axis of variation if data are normalized by leaf mass. We show that these traits are approximately distributed proportional to leaf area instead of mass, as expected for a light- and carbon dioxide-collecting organ. Much of the structure in the mass-normalized LES results from normalizing area-proportional traits by mass. Mass normalization induces strong correlations among area-proportional traits because of large variation among species in leaf mass per area (LMA). The high LMA variance likely reflects its functional relationship with leaf life span. A LES that is independent of mass- or area-normalization and LMA reveals physiological relationships that are inconsistent with those in global vegetation models designed to address climate change.