Projected changes in temperature and drought regime are likely to reduce carbon (C) storage in forests, thereby amplifying rates of climate change. While such reductions are often presumed to be greatest in semi-arid forests that experience widespread tree mortality, the consequences of drought may also be important in temperate mesic forests of Eastern North America (ENA) if tree growth is significantly curtailed by drought. Investigations of the environmental conditions that determine drought sensitivity are critically needed to accurately predict ecosystem feedbacks to climate change. We matched site factors with the growth responses to drought of 10,753 trees across mesic forests of ENA, representing 24 species and 346 stands, to determine the broad-scale drivers of drought sensitivity for the dominant trees in ENA. Here we show that two factors-the timing of drought, and the atmospheric demand for water (i.e., local potential evapotranspiration; PET)-are stronger drivers of drought sensitivity than soil and stand characteristics. Drought-induced reductions in tree growth were greatest when the droughts occurred during early-season peaks in radial growth, especially for trees growing in the warmest, driest regions (i.e., highest PET). Further, mean species trait values (rooting depth and ψ ) were poor predictors of drought sensitivity, as intraspecific variation in sensitivity was equal to or greater than interspecific variation in 17 of 24 species. From a general circulation model ensemble, we find that future increases in early-season PET may exacerbate these effects, and potentially offset gains in C uptake and storage in ENA owing to other global change factors.

In fall (November 2005) and winter (February 2006), we collected current-year foliage of native red spruce (Picea rubens Sarg.) growing in a reference watershed and in a watershed treated in 1999 with wollastonite (CaSiO(3), a slow-release calcium source) to simulate preindustrial soil calcium concentrations (Ca-addition watershed) at the Hubbard Brook Experimental Forest (Thornton, NH). We analyzed nutrition, soluble sugar concentrations, ascorbate peroxidase (APX) activity and cold tolerance, to evaluate the basis of recent (2003) differences between watersheds in red spruce foliar winter injury. Foliar Ca and total sugar concentrations were significantly higher in trees in the Ca-addition watershed than in trees in the reference watershed during both fall (P=0.037 and 0.035, respectively) and winter (P=0.055 and 0.036, respectively). The Ca-addition treatment significantly increased foliar fructose and glucose concentrations in November (P=0.013 and 0.007, respectively) and foliar sucrose concentrations in winter (P=0.040). Foliar APX activity was similar in trees in both watersheds during fall (P=0.28), but higher in trees in the Ca-addition watershed during winter (P=0.063). Cold tolerance of foliage was significantly greater in trees in the Ca-addition watershed than in trees in the reference watershed (P<0.001). Our results suggest that low foliar sugar concentrations and APX activity, and reduced cold tolerance in trees in the reference watershed contributed to their high vulnerability to winter injury in 2003. Because the reference watershed reflects forest conditions in the region, the consequences of impaired physiological function caused by soil Ca depletion may have widespread implications for forest health.

No abstract

Forests of northeastern North America have been exposed to anthropogenic acidic inputs for decades, resulting in altered cation relations and disruptions to associated physiological processes in multiple tree species, including sugar maple (Acer saccharum Marsh.). In the current study, the impacts of calcium (Ca) and aluminum (Al) additions on mature sugar maple physiology were evaluated at the Hubbard Brook Experimental Forest (Thornton, NH, USA) to assess remediation (Ca addition) or exacerbation (Al addition) of current acidified conditions. Fine root cation concentrations and membrane integrity, carbon (C) allocation, foliar cation concentrations and antioxidant activity, foliar response to a spring freezing event and reproductive ability (flowering, seed quantity, filled seed and seed germination) were evaluated for dominant sugar maple trees in a replicated plot study. Root damage and foliar antioxidant activity were highest in Al-treated trees, while growth-associated C, foliar re-flush following a spring frost and reproductive ability were highest in Ca-treated trees. In general, we found that trees on Ca-treated plots preferentially used C resources for growth and reproductive processes, whereas Al-treated trees devoted C to defense-based processes. Similarities between Al-treated and control trees were observed for foliar cation concentrations, C partitioning and seed production, suggesting that sugar maples growing in native forests may be more stressed than previously perceived. Our experiment suggests that disruption of the balance of Ca and Al in sugar maples by acid deposition continues to be an important driver of tree health.

Red spruce (Picea rubens Sarg.) trees are uniquely vulnerable to foliar freezing injury during the cold season (fall and winter), but are also capable of photosynthetic activity if temperatures moderate. To evaluate the influence of calcium (Ca) addition on the physiology of red spruce during the cold season, we measured concentrations of foliar polyamines and free amino acids (putative stressprotection compounds), chlorophyll (a key photosystem component), and sapwood area (a proxy for foliar biomass), for trees in Ca-addition (CaSiO 3 added) and Cadepleted (reference) watersheds at the Hubbard Brook Experimental Forest (NH, USA). Ca-addition increased concentrations of the amino acids alanine and c-aminobutyric acid (GABA) and the polyamines putrescine (Put) and spermidine (Spd) in November, and Put in February relative to foliage from the reference watershed. Consistent with increased stress protection, foliage from the Caaddition watershed had higher total chlorophyll and chlorophyll a concentrations in February than foliage from the reference watershed. In contrast, foliage from the reference watershed had significantly lower glutamic acid (Glu) and higher alanine (Ala) concentrations in February than foliage from the Ca-addition watershed. Imbalances in Ala:Glu have been attributed to cold sensitivity or damage in other species. In addition to concentration-based differences in foliar compounds, trees from the Ca-addition watershed had higher estimated levels of foliar biomass than trees from the reference watershed. Our findings suggest that Caaddition increased the stress tolerance and productive capacity of red spruce foliage during the cold season, and resulted in greater crown mass compared to trees growing on untreated soils.