Background and Aims

Trees universally decrease their growth with age. Most explanations for this trend so far support the hypothesis that carbon (C) gain becomes limited with age; though very few studies have directly assessed the relative reductions of C gain and C demand with tree age. It has also been suggested that drought enhances the effect of C gain limitation in trees. Here tests were carried out to determine whether C gain limitation is causing the growth decay with tree age, and whether drought accentuates its effect.

Methods

The balance between C gain and C demand across tree age and height ranges was estimated. For this, the concentration of non-structural carbohydrates (NSCs) in stems and roots of trees of different ages and heights was measured in the deciduous temperate species Nothofagus pumilio. An ontogenetic decrease in NSCs indicates support for C limitation. Furthermore, the importance of drought in altering the C balance with ontogeny was assessed by sampling the same species in Mediterranean and humid climate locations in the southern Andes of Chile. Wood density (WD) and stable carbon isotope ratios (δ13C) were also determined to examine drought constraints on C gain.

Key Results

At both locations, it was effectively found that tree growth ultimately decreased with tree age and height. It was found, however, that NSC concentrations did not decrease with tree age or height when WD was considered, suggesting that C limitation is not the ultimate mechanism causing the age/height-related declining tree growth. δ13C decreased with tree age/height at the Mediterranean site only; drought effect increased with tree age/height, but this pattern was not mirrored by the levels of NSCs.

Conclusions

The results indicate that concentrations of C storage in N. pumilio trees do not decrease with tree age or height, and that reduced C assimilation due to summer drought does not alter this pattern.

Background and Aims

A long-running debate centres on whether shade tolerance of tree seedlings is mainly a function of traits maximizing net carbon gain in low light, or of traits minimizing carbon loss. To test these alternatives, leaf display, light-interception efficiency, and simulated net daily carbon gain of juvenile temperate evergreens of differing shade tolerance were measured, and how these variables are influenced by ontogeny was queried.

Methods

The biomass distribution of juveniles (17–740 mm tall) of seven temperate rainforest evergreens growing in low (approx. 4 %) light in the understorey of a second-growth stand was quantified. Daytime and night-time gas exchange rates of leaves were also determined, and crown architecture was recorded digitally. YPLANT was used to model light interception and carbon gain.

Results

An index of species shade tolerance correlated closely with photosynthetic capacities and respiration rates per unit mass of leaves, but only weakly with respiration per unit area. Accumulation of many leaf cohorts by shade-tolerant species meant that their ratios of foliage area to biomass (LAR) decreased more gradually with ontogeny than those of light-demanders, but also increased self-shading; this depressed the foliage silhouette-to-area ratio (STAR), which was used as an index of light-interception efficiency. As a result, displayed leaf area ratio (LARd = LAR × STAR) of large seedlings was not related to species shade tolerance. Self-shading also caused simulated net daily carbon assimilation rates of shade-tolerant species to decrease with ontogeny, leading to a negative correlation of shade tolerance with net daily carbon gain of large (500 mm tall) seedlings in the understorey.

Conclusions

The results suggest that efficiency of energy capture is not an important correlate of shade tolerance in temperate rainforest evergreens. Ontogenetic increases in self-shading largely nullify the potential carbon gain advantages expected to result from low respiration rates and long leaf lifespans in shade-tolerant evergreens. The main advantage of their long-lived leaves is probably in reducing the costs of crown maintenance.