The effects of defoliation on carbon allocation: can carbon limitation reduce growth in favour of storage?

There is no consensus about how stresses such as low water availability and temperature limit tree growth. Sink limitation to growth and survival is often inferred if a given stress does not cause non-structural carbohydrate (NSC) concentrations or levels to decline along with growth. However, trees may actively maintain or increase NSC levels under moderate carbon stress, making the pattern of reduced growth and increased NSCs compatible with carbon limitation. To test this possibility, we used full and half defoliation to impose severe and moderate carbon limitation on 2-year-old Quercus velutina Lam. saplings grown in a common garden. Saplings were harvested at either 3 weeks or 4 months after treatments were applied, representing short- and longer-term effects on woody growth and NSC levels. Both defoliation treatments maintained a lower total leaf area than controls throughout the experiment with no evidence of photosynthetic up-regulation, and resulted in a similar total biomass reduction. While fully defoliated saplings had lower starch levels than controls in the short term, half defoliated saplings maintained control starch levels in both the short and longer term. In the longer term, fully defoliated saplings had the greatest starch concentration increment, allowing them to recover to near-control starch levels. Furthermore, between the two harvest dates, fully and half defoliated saplings allocated a greater proportion of new biomass to starch than did controls. The maintenance of control starch levels in half defoliated saplings indicates that these trees actively store a substantial amount of carbon before growth is carbon saturated. In addition, the allocation shift favouring storage in defoliated saplings is consistent with the hypothesis that, as an adaptation to increasing carbon stress, trees can prioritize carbon reserve formation at the expense of growth. Our results suggest that as carbon limitation increases, reduced growth is not necessarily accompanied by a decline in NSC concentrations. Therefore, a lack of NSC decline may not be evidence that reduced tree growth under cold or water stress is caused by sink limitation.