Relative growth rate in relation to physiological and morphological traits for northern hardwood tree seedlings: species, light environment and ontogenetic considerations.

The influence of ontogeny, light environment and species on relationships of relative growth rate (RGR) to physiological and morphological traits were examined for first-year northern hardwood tree seedlings. Three Betulaceae species (Betula papyrifera, Betula alleghaniensis and Ostrya virginiana) were grown in high and low light and Quercus rubra and Acer saccharum were grown only in high light. Plant traits were determined at four ages: 41, 62, 83 and 104 days after germination. In high light (610 μmol m-2 s-1 PPFD), across species and ages, RGR was positively related to the proportion of the plant in leaves (leaf weight ratio, LWR; leaf area ratio, LAR), in situ rates of average canopy net photosynthesis (A) per unit mass (Amass) and per unit area (Aarea), and rates of leaf, stem and root respiration. In low light (127 μmol m-2 s-1 PPFD), RGR was not correlated with Amass and Aarea whereas RGR was positively correlated with LAR, LWR, and rates of root and stem respiration. RGR was negatively correlated with leaf mass per area in both high and low light. Across light levels, relationships of CO2 exchange and morphological characteristics with RGR were generally weaker than within light environments. Moreover, relationships were weaker for plant parameters containing a leaf area component (leaf mass per area, LAR and Aarea), than those that were solely mass-based (respiration rates, LWR and Amass). Across light environments, parameters incorporating the proportion of the plant in leaves and rates of photosynthesis explained a greater amount of variation in RGR (e.g. LWR*Amass, R2=0.64) than did any single parameter related to whole-plant carbon gain. RGR generally declined with age and mass, which were used as scalars of ontogeny. LWR (and LAR) also declined for seven of the eight species-light treatments and A declined in four of the five species in high light. Decreasing LWR and A with ontogeny may have been partially responsible for decreasing RGR. Declines in RGR were not due to increased respiration resulting from an increase in the proportion of solely respiring tissue (roots and stems). In general, although LWR declined with ontogeny, specific rates of leaf, stem, and root respiration also decreased. The net result was that whole-plant respiration rates per unit leaf mass decreased for all eight treatments. Identifying the major determinants of variation in growth (e.g. LWR*Amass) across light environments, species and ontogeny contributes to the establishment of a framework for exploring limits to productivity and the nature of ecological success as measured by growth. The generality of these relationships both across the sources of variation we explored here and across other sources of variation in RGR needs further study.