Increased Photosynthesis Offsets Costs of Allocation to Sapwood in An Arid Environment

We assessed the effect that varying patterns of biomass allocation had on growth of ponderosa pine (Pinus ponderosa) growing in the desert climate of the Great Basin and the montane climate of the eastern Sierra Nevada. Prior work established that desert trees have lower leaf: sapwood area ratios than montane trees (0.104 and 0.201 M2/ cm2, respectively) and proportionally greater stem respiration. Sapwood: leaf mass ratios are also greater and increase more as a function stem diameter in desert than in montane trees. We hypothesized that this increased allocation of carbon to stem sapwood and stem respiration in large trees could decrease growth rates in the desert compared to the montane environment, in addition to any growth reduction imposed by drought on physiology and growth processes. Trees of all diameters (dbh) in the desert environment had lower relative growth rates (RGRs) than montane trees (e.g., for a 30 cm dbh tree, RGR = 0.012 vs. 0.021 kg-kg-1-yr-', respectively). However, growth rates of desert and montane trees declined similarly with increasing dbh and did not reflect diverging sapwood: leaf mass ratios. Alternatively, we hypothesized that desert trees may increase rates of photosynthetic carbon accumulation (per unit leaf area) with diameter, thereby compensating for increased sapwood respiration. Leaf nitrogen (N) concentration and stable-carbon isotope composition (813C) were measured to examine size-dependent and seasonally integrated photosynthetic capacity within desert and montane environments. Nitrogen concentration was correlated with photosynthetic capacity. Leaf nitrogen (N) concentration and b'3C values were greater in the desert (e.g., in 1-yr-old needles, desert 1.11% and -22.96%o; montane = 0.94% and -24.20%o) and differed between desert and montane trees as a function of dbh. In desert trees, leaf nitrogen concentration in 1-yr-old through 5-yr-old needles increased with dbh, and carbon isotope composition in 1-yr-old needles increased with dbh, suggesting increased photosynthetic capacity and photosynthetic rates with increasing tree size. Needle nitrogen concentration and b83C values decreased or remained constant with dbh in montane trees. Desert trees had greater light extinction coefficients and retained fewer needle cohorts. Our results suggest that increased allocation to heterotrophic stem tissue at the expense of photosynthetic tissue does not always incur a reduction in tree growth as predicted by current models of forest productivity.