RESUMO
There is an ongoing debate on whether a drought induced carbohydrate limitation (source limitation) or a direct effect of water shortage (sink limitation) limit growth under drought. In this study, we investigated the effects of the two driest summers recorded in southern Chile in the last seven decades, on the growth and non-structural carbohydrates (NSC) concentrations of the slow-growing conifer Fitzroya cupressoides. Specifically, we studied the seasonal variation of NSC in saplings and adults one and two years after the occurrence of a 2 year-summer drought at two sites of contrasting precipitation and productivity (mesic-productive vs. rainy-less productive). We also evaluated radial growth before, during and after the drought, and predicted that drought could have reduced growth. If drought caused C source limitation, we expected that NSCs will be lower during the first than the second year after drought. Conversely, similar NSC concentrations between years or higher NSC concentrations in the first year would be supportive of sink limitation. Also, due to the lower biomass of saplings compared with adults, we expected that saplings should experience stronger seasonal NSC remobilization than adults. We confirmed this last expectation. Moreover, we found no significant growth reduction during drought in the rainy site and a slightly significant growth reduction at the mesic site for both saplings and adults. Across organs and in both sites and age classes, NSC, starch, and sugar concentrations were generally higher in the first than in the second year following drought, while NSC seasonal remobilization was generally lower. Higher NSC concentrations along with lower seasonal NSC remobilization during the first post-drought year are supportive of sink limitation. However, as these results were found at both sites while growth decreased slightly and just at the mesic site, limited growth only is unlikely to have caused NSC accumulation. Rather, these results suggest that the post-drought dynamics of carbohydrate storage are partly decoupled from the growth dynamics, and that the rebuild of C reserves after drought may be a priority in this species.
RESUMO
Seedlings of nine southern Chilean trees were grown at three nutrient supply rates, to examine the roles of growth rate, biomass distribution and nutrient use traits in determining species natural distributions on resource gradients. Relative growth rate (RGR) showed no overall relationship with species site requirements, although RGR of fertile-site species tended to be more responsive to nutrient supply. In the low-nutrient treatment, infertility-tolerant Fitzroya cupressoides showed a higher RGR rank than a fertility-demanding species (Laurelia philippiana) which outgrew it substantially at the highest supply rate. This reversal of RGR ranks was associated with divergent nutrient use responses: at high nutrient supply both spp. had similar plant nitrogen concentrations (PNC), whereas at the low supply rate Fitzroya's production of biomass per unit of assimilated N was twice that of Laurelia's. However, this pattern does not appear to serve as a general explanation of the respective distributions of the study species, as RGR ranks of most species were unaltered by nutrient supply. At low nutrient availability, no clear differences in shoot:root ratio (SRR) were apparent between poor-site and fertile-site species. However, at high nutrient availability, SRR was markedly higher in the latter, resulting from differences in biomass allocation to stems (not leaves). Leaf area ratios (LAR) were higher in fertile-site species than in those tolerant of low fertility, because of differences in specific leaf area rather than leaf weight ratio. Very high LAR at high nutrient supply was characteristic of most shade-tolerant angiosperms, but not of shade-tolerant conifers. Although PNC showed no overall differences between poor- and fertile-site species, sensitivity of PNC to external supply rate was greatest in two infertility-tolerant conifers. In contrast, the angiosperm Weinmannia trichosperma, although tolerant of low fertility, responded to increased nutrient supply with greatly increased RGR and little change in PNC. Results show little trait convergence between conifers and angiosperms in adaptation both to shade and to infertile soils; i.e. fitness of different taxa in a given environment may hinge on different trait combinations.