Gary J. Hawley

Acid Rain Impacts on Calcium Nutrition and Forest Health

Forest ecosystems throughout the world are exposed to acid rain, a complex solution consisting largely of H+, SO42-, NH4+, and NO3- pollutant ions derived from sulfur and nitrogen oxides. Although the public in the United States may view acid deposition as a localized pollution issue specific to certain regions of North America and northern Europe, recent evidence of acidic deposition throughout much of Asia, including tropical forest regions, has demonstrated that acid deposition is actually a global phenomenon (Abate 1995).

Association of red coloration with senescence of sugar maple leaves in autumn

We evaluated the association of red coloration with senescence in sugar maple (Acer saccharum Marsh.) leaves by assessing differences in leaf retention strength and the progression of the abscission layer through the vascular bundle of green, yellow, and red leaves of 14 mature open-grown trees in October 2002. Computer image analysis confirmed visual categorization of leaves as predominantly green, yellow or red, and chemical quantification of leaf pigment concentrations verified that leaf color reflected underlying differences in leaf biochemistry. Significantly lower chlorophyll concentrations within red and yellow leaves indicated that senescence was more advanced in leaves from these color categories relative to green leaves. Among leaf types, only red leaves contained high concentrations of anthocyanins. There were significant differences in leaf retention capacity among color categories, with the petioles of green leaves being the most firmly attached to twigs, followed by red and then yellow leaves. Microscopic analysis indicated that yellow leaves had the most advanced extension of the abscission layer through the vasculature, with green and red leaves having significantly less abscission layer progression than yellow. A more limited progression of the abscission layer through vascular bundles may be evidence of delayed leaf senescence that could extend resorption of mobile leaf constituents. Together, results from this study suggest an association between leaf anthocyanin content and functional delays in senescence.

Calcium and aluminum impacts on sugar maple physiology in a northern hardwood forest

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.

Calcium addition at the Hubbard Brook Experimental Forest increases the capacity for stress tolerance and carbon capture in red spruce (Picea rubens) trees during the cold season

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 stress-protection compounds), chlorophyll (a key photosystem component), and sapwood area (a proxy for foliar biomass), for trees in Ca-addition (CaSiO3 added) and Ca-depleted (reference) watersheds at the Hubbard Brook Experimental Forest (NH, USA). Ca-addition increased concentrations of the amino acids alanine and γ-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 Ca-addition 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 Ca-addition 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.

Assessment of weather-associated causes of red spruce winter injury and consequences to aboveground carbon sequestration

Despite considerable study, it remains uncertain what environmental factors contribute to red spruce (Picea rubens Sarg.) foliar winter injury and how much this injury influences tree C stores. We used a long-term record of winter injury in a plantation in New Hampshire and conducted stepwise linear regression analyses with local weather and regional pollution data to determine which parameters helped account for observed injury. Two types of weather phenomena were consistently associated with elevated injury: (i) measures of low-temperature stress that incite injury and (ii) factors that reduced the length of the growing season and predisposed trees to injury. At this plantation, there was a significant linear relationship between winter injury and growth reductions for 2 years after a severe winter injury event. Analysis using data from three New England states indicated that plantation data reflected a regional response.

Effects of soil calcium and aluminum on the physiology of balsam fir and red spruce saplings in northern New England

We examined the influence of calcium (Ca) and aluminum (Al) nutrition on the foliar physiology of red spruce (Picea rubens Sarg.) and balsam fir [Abies balsamea (L.) Mill.] in northern New England, USA. At the Hubbard Brook Experimental Forest (NH, USA), spruce and fir saplings were sampled from control, Al-, and Ca-supplemented plots at a long-established nutrient perturbation (NuPert) study in fall 2008. Measurements included cation concentrations (roots and foliage), dark-adapted chlorophyll fluorescence (Fv/Fm), soluble sugar concentrations, and ascorbate peroxidase (APX) and glutathione reductase (GR) activity in current-year foliage. Additional untreated saplings were sampled from base-rich Sleepers River (VT) and base-poor Jeffers Brook (NH) for Fv/Fm and foliar nutrient concentrations. At NuPert, there were significantly greater Ca concentrations and Ca:Al ratios in roots from the Ca end vs. the Al end of the Al-control-Ca addition gradient. There were also trends toward greater foliar Ca and Ca:Al ratios and lower Al concentrations across the treatment gradient at NuPert and for foliage at Sleepers River vs. Jeffers Brook. At NuPert, Fv/Fm and APX activity increased across the treatment gradient, and red spruce was higher in these measures than balsam fir. These patterns were also observed when Jeffers Brook and Sleepers River were compared. Increased Ca availability appeared to enhance the ability of red spruce and balsam fir to repair oxidative stress damage, including photooxidation. Our findings support work indicating a greater contemporary level of stress for balsam fir relative to red spruce, which is surprising considering the well-documented regional decline of spruce.

Is nut cold tolerance a limitation to the restoration of American chestnut in the northeastern United States

American chestnut (Castanea dentata) was once a dominant hardwood species in the eastern United States, growing from Maine to Georgia and west to the Ohio Valley (Harlow et al. 1979). Arguably, American chestnut may have been the most important hardwood species in North America, renowned for its quick growth, massive size, and great utility (Harlow et al. 1979). Unfortunately, within 50 years of the introduction of chestnut blight (Cryphonectria parasitica)--a fungal disease native to Asia--American chestnut was functionally removed as an overstory tree from eastern forests (Griffin 2000).

Inconclusive evidence of Juniperus virginiana recovery following sulfur pollution reductions

Thomas et al. (1) address a question of great scientific interest: have pollution reductions mandated by the Clean Air Act improved forest health and productivity? Although answers to this question are of great importance, various aspects of this work limit its ability to address this question.

The surprising recovery of red spruce growth shows links to decreased acid deposition and elevated temperature

Following growth declines and increased mortality linked to acid deposition-induced calcium depletion, red spruce (Picea rubens Sarg.) in the northeastern United States are experiencing a recovery. We found that more than 75% of red spruce trees and 90% of the plots examined in this study exhibited increasing growth since 2001. To understand this change, we assessed the relationship between red spruce radial growth and factors that may influence growth: tree age and diameter, stand dynamics, plot characteristics (elevation, slope, aspect, geographical position), and a suite of environmental variables (temperature, precipitation, climate and precipitation indices (degree days, SPEI [standardized precipitation evapotranspiration index], and acid deposition [SO42-, NO3-, pH of rainfall, cation:anion ratio of rainfall]) for 52 plots (658 trees) from five states (spanning 2.5°N × 5°W). Examining the growth relationships from 1925 to 2012, we found that while there was variability in response to climate and acid deposition (limited to 1980-2012) by elevation and location, plot and tree factors did not adequately explain growth. Higher temperatures outside the traditional growing season (e.g., fall, winter, and spring) were related to increased growth. Nitrogen deposition (1980-2012) was associated with lower growth, but the strength of this relationship has lessened over time. Overall, we predict sustained favorable conditions for red spruce in the near term as acid deposition continues to decline and non-traditional growing season (fall through spring) temperatures moderate, provided that overall temperatures and precipitation remain adequate for growth.

Quantifying Ecosystem Controls and Their Contextual Interactions on Nutrient Export from Developing Forest Mesocosms

The complexity of natural ecosystems makes it difficult to compare the relative importance of abiotic and biotic factors and to assess the effects of their interactions on ecosystem development. To improve our understanding of ecosystem complexity, we initiated an experiment designed to quantify the main effects and interactions of several factors that are thought to affect nutrient export from developing forest ecosystems. Using a replicated 2 × 2 × 4 factorial experiment, we quantified the main effects of these factors and the factor interactions on annual calcium, magnesium, and potassium export from field mesocosms over 4 years for two Vermont locations, two soils, and four different tree seedling communities. We found that the main effects explained 56%–97% of total variation in nutrient export. Abiotic factors (location and soil) accounted for a greater percentage of the total variation in nutrient export (47%–94%) than the biotic factor (plant community) (2%–15%). However, biotic control over nutrient export was significant, even when biomass was minimal. Factor interactions were often significant, but they explained less of the variation in nutrient export (1%–33%) than the main effects. Year-to-year fluctuations influenced the relative importance of the main effects in determining nutrient export and created factor interactions between most of the explanatory variables. Our study suggests that when research is focused on typically used main effects, such as location and soil, and interactions are aggregated into overall error terms, important information about the factors controlling ecosystem processes can be lost.