Source: UNIVERSITY OF VERMONT submitted to NRP
THE ROLE OF CA DEPLETION AND WINTER INJURY ON THE PRODUCTIVITY OF RED SPRUCE IN VERMONT
Sponsoring Institution
National Institute of Food and Agriculture
Project Status
COMPLETE
Funding Source
Reporting Frequency
Annual
Accession No.
0208593
Grant No.
(N/A)
Cumulative Award Amt.
(N/A)
Proposal No.
(N/A)
Multistate No.
(N/A)
Project Start Date
Oct 1, 2006
Project End Date
Sep 30, 2010
Grant Year
(N/A)
Program Code
[(N/A)]- (N/A)
Recipient Organization
UNIVERSITY OF VERMONT
(N/A)
BURLINGTON,VT 05405
Performing Department
SCHOOL OF NATURAL RESOURCES
Non Technical Summary
Even though Ca depletion may increase the risk of red spruce winter injury, it is not clear how this translates to changes in growth and productivity for trees in the field. Earlier work demonstrated that moderate winter injury can reduce the woody growth of red spruce. However, there has been no simultaneous evaluation of growth impacts from a broad range of injury levels, including severe injury that may ultimately lead to stand decline. In this study we will conduct assessments radial tree growth and foliar Ca nutrition to quantitatively integrate and model the influence of Ca depletion on winter injury and productivity of red spruce trees across the region. Resolution of this issue is needed by scientists, managers and policy makers in order to complete a more comprehensive analysis of the biological and economic consequences of acid rain-induced Ca depletion.
Animal Health Component
40%
Research Effort Categories
Basic
60%
Applied
40%
Developmental
(N/A)
Classification

Knowledge Area (KA)Subject of Investigation (SOI)Field of Science (FOS)Percent
1020610102050%
1330430101030%
2030610102010%
2050610310010%
Goals / Objectives
Our overall objective for this research is to determine if anthropogenic Ca deletion has the potential to deplete biological Ca reserves and predispose trees to exaggerated injury, reduced growth and decline following environmental stress exposures. More specifically we will quantitatively evaluate the influence of foliar winter injury on the productivity of red spruce trees. Because winter injury is predisposed by deficiencies in biological Ca reserves this project will specifically address how Ca depletion influences winter injury and growth reductions that can lead to stand decline. One of our objective is to use data generated in this project to estimate overall reductions in xylem growth and biomass accumulation that result from winter injury, enabling us to quantitatively evaluate the links between Ca nutrition, foliar injury, and reductions in forest productivity. Our objetive is to utilize a multi-scale approach involving assessments of individual trees, stand-level evaluations, and modeling that expands the relevance of results to the Northern Forest region.
Project Methods
Recent preliminary data provides strong evidence that red spruce foliar winter injury can result in significant (up to 50%) and extended growth reductions, and suggests that the relationship between winter injury and growth can be mathematically modeled to quantify the impacts of damage. We propose a multi-phase study to build upon this preliminary data and specifically quantify 1) the impact of winter injury on red spruce growth and biomass accumulation, and 2) the influence of Ca depletion on injury-associated growth declines. In phase one of this study we will refine analysis of preliminary data of winter injury and growth of individual trees to develop a mathematical model of the relationship between the amount of winter injury in 2003 (the most recent and severe episode of region-wide injury) and subsequent reductions of xylem growth. Information on the impacts of winter injury on growth and biomass measures will provide unique and needed insight into the ecological and potential economic impacts of winter injury. We will field-verify the mathematical model refined in phase 1 by assessing mean stand-level growth of trees growing at a sub-set of 22 sites across Vermont where winter injury was assessed in 2003 (Lazarus et al. 2004). Two sites with mean winter injury damage levels representing each of 11 injury categories (0% damage to current-year foliage, 1-10% damage to current-year foliage , etc. up to 100% damage) will be assessed to evaluate the full range of injury and potential impacts on growth. We will utilize our model of growth reductions from winter injury, in conjunction with previous data on the spatial distribution and intensity of foliar winter injury (Lazarus et al. 2006), to estimate growth reductions resulting from winter injury in 2003 for the broader Northern Forest region. A model developed during a recent project highlighted specific geographic factors that affected the extent and severity of 2003 winter injury, including elevation, longitude, and plot aspect (R2 = 0.83; Lazarus et al. 2006). Trees at high elevation in the western part of the range and on west facing slopes receive higher levels of acid loading and are predicted to have lower levels of available Ca. Our proposed project will utilize that model, and allow us to quantitatively estimate variations in growth reductions across the region. We will assess the role of Ca availability in mitigating winter injury-related growth declines of trees growing on Ca-fertilized and unfertilized sites. We will assess mean site-level growth in plots in which we measured winter injury in 2003. Growth data from these sites will be compared with estimates generated by the model created and refined during previous phases of this project to determine whether Ca fertilization can mitigate growth reductions associated with observed levels of winter injury, or whether Ca may impart additional growth benefits beyond what is expected from our winter injury-growth model. This phase will allow us to more directly link the contribution of Ca nutrition to changes in winter injury and reductions in growth and biomass production.

Progress 10/01/06 to 09/30/10

Outputs
OUTPUTS: Field collections and assessments conducted throughout this project were designed to evaluate the influence of winter injury and calcium (Ca) nutrition on Ca dependent physiology including growth and carbon (C) sequestration of red spruce. Using foliage collected from red spruce growing in a Ca depleted reference watershed and in a watershed treated in 1999 with wollastonite (slow-release Ca source) at the Hubbard Brook Experimental Forest (NH), the nutrition, concentrations of stress related compounds, photosystem function, and sapwood area, were analyzed to assess the influence of Ca addition on the protective biochemistry and C capture capacity of foliage during the cold season. Analysis of the Ca concentrations of a subsample of foliage from the same collections showed higher concentrations in trees in the Ca-addition than in foliage from trees in the reference watershed. Coincident with differences in Ca nutrition, the amino acids alanine (Ala) and I3-aminobutyric acid (GABA), and the polyamines putrescine (Put) and spermidine (Spd) concentrations were different in trees in the Ca-addition compared to the reference watershed. These polyamines and GABA are thought to have direct protective or other benefits to leaves that may contribute to stress tolerance. The foliage of trees from the Ca-addition had higher total chlorophyll and chlorophyll a concentrations than foliage in trees in the reference watershed in Feb - indicating that Ca-supplemented trees likely retained a greater capacity for winter photosynthesis. Trees from the Ca-addition watershed also had higher estimated levels of overall foliar biomass than trees from the reference watershed. Together, our findings indicate that Ca-addition resulted in more and better-protected red spruce foliage that had an enhanced capacity for winter C capture compared to the foliage of trees that were subject to ambient soil Ca concentrations. We assessed the causes for year-to-year variations in red spruce foliar winter injury and the influence of injury on xylem growth and carbon (C) sequestration. We used a long-term record of winter injury in a plantation in NH. 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. There was a linear relationship between winter injury and growth reductions for two years after a severe winter injury event. Analysis using data from three New England states indicated that plantation data reflected a regional response. Using regional data we estimated a reduction of 394,000 metric tons of C sequestered in red spruce stems greater than 20 cm in diameter growing in NY and northern New England during the two years following a severe winter injury event. Reductions in C sequestration likely alter nutrient and water cycling, and other ecosystem services that are influenced by C uptake. PARTICIPANTS: Dr. Paul G. Schaberg and his research team at the Burlington Laboratory of the U.S. Forest Service were major participants in all aspects of this research. We have also worded closely with Dr. C. Eagar of the US Forest Service in Durham, NH as well as numerous researcher (Harvard, Cornell, Institute of Ecosystem Studies, Boston College) working at Hubbard Brook Experimental Forest in Thronton, NH. There were numerous opportunities for training and professional development for several undergraduate students, Masters students and PhD students conducting research associated with this project. TARGET AUDIENCES: Land managers in the northeastern United States and Canada benefit from knowledge pertaining to the implications of continued pollution inputs into forested ecosystems that result in nutrient depletions. In addition, policy makers also benefit from data summarizing the implication of acid-rain induced calcium depletion on our northern forests. Other scientists will also utilize our results to aid them in understanding the implication of calcium depletion including modelers that use such data to predict growth and product losses associated with anthropogenic inputs. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
Results from this project provide the first quantitative evaluation of how pollution-influenced winter injury affects the health and productivity of red spruce, and enables us to quantitatively evaluate the links between Ca nutrition, foliar injury, and changes in forest productivity. This work provides contextual information for policy makers and managers seeking to understand the potential landscape-scale impacts of continued acidic deposition and Ca depletion, and provides data needed for possible future work to evaluate the economic impacts of pollution-exacerbated winter injury. This work (combined with past findings) will help evaluate the cause(s) for the decline of an economically and ecologically important tree species, and help assess the possibility that Ca depletion more broadly predisposes trees (red spruce and other important species to the northeastern US and southeastern Canada such as sugar maple and paper birch) to inadequate stress response and decline - a scenario with significant management and policy implications for the Northern Forest.

Publications

  • Schaberg, P.G.; Murakami, P.F.; Turner, M.R.; Heitz, H.K.; Hawley, G.J. 2008. Association of red coloration with senescence of sugar maple leaves in autumn. Trees. 22: 573-578.
  • Murakami, P.F.; Schaberg, P.G.; Shane, J.B. 2008. Stem girdling manipulates leaf sugar concentrations and anthocyanin expression in sugar maples trees during autumn. Tree Physiology. 28: 1467-1473.
  • Strimbeck, G. Richard; Kjellsen, Trygve D.; Schaberg, Paul G.; Murakami, Paula F. 2007. Cold in the common garden: comparative low-temperature tolerance of boreal and temperate conifer foliage. Trees. 21: 557-567.
  • Schaberg, P.G.; Hawley, G.J.; Halman, J.M.; Murakami, P.F. 2009. Laboratory and field data indicate that acidic deposition-induced calcium depletion disrupts the nutrition and physiology of trees, predisposing them to decline. Proceedings of the National Acidic Deposition Program Annual Meeting and Scientific Symposium, Saratoga Springs, New York, October 7, 2009, p44.
  • Schaberg, P.G.; Hawley, G.J.; Elliott, H.K.; Murakami, P.F.; Huggett, B.A.; Halman, J.M.; Eagar, C. 2009. Long-term fertilization with calcium or aluminum highlights the influence of anthropogenic cation disruption on the physiology and carbon sequestration of sugar maple trees at the Hubbard Brook Experimental Forest, NH. 94th annual meeting of the Ecological Society of America; August 2-7; Albuquerque, NM: Ecological Society of America. Available at http://eco.confex.com/eco/2009/techprogram/P18570.HTM. Abstract.
  • Halman, J.M.; Schaberg, P.G.; Hawley, G.J.; Hansen, C.F. 2009. Potential role of Ca in the response of paper birch (Betula papyrifera) to ice storm-induced decline in Vermont, USA. In: 94th annual meeting of the Ecological Society of America; 2009 August 2-7; Albuquerque, NM: Ecological Society of America. Available at http://eco.confex.com/eco/2009/techprogram/P18378.HTM. Abstract.
  • Pardo, L.H.; Semaoune, P.; Schaberg, P.G.; Eagar C.; Sebilo, M. 2009. Patterns in I'15N in roots, stems and leaves of sugar maple and beech seedlings, saplings, and mature trees. In: 94th annual meeting of the Ecological Society of America; 2009 August 2-7; Albuquerque, NM: Ecological Society of America. Available at http://eco.confex.com/eco/2009/techprogram/P16777.HTM. Abstract.
  • Schaberg, Paul G.; Hawley, Gary J. 2010. Disruption of calcium nutrition at Hubbard Brook Experimental Forest (New Hampshire) alters the health and productivity of red spruce and sugar maple trees and provides lessons pertinent to other sites and regions. In: Rentch, James S.; Schuler, Thomas M., eds. 2010. Proceedings conference on the ecology and management of high-elevation forests in the central and southern Appalachian Mountains. 2009 May 14-15; Slatyfork, WV. Gen. Tech. Rep. NRS-P-64. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 190-200.
  • Strimbeck, G.R.; Schaberg, P.G. 2009. Going to extremes: low temperature tolerance and acclimation in temperate and boreal conifers. In: Gusta L.; Wisniewski, M.; Tanino, K., eds. Plant cold hardiness: from the laboratory to the field. CAB International: 226-239.


Progress 10/01/08 to 09/30/09

Outputs
OUTPUTS: Using foliage collected from red spruce growing in a calcium (Ca) depleted reference watershed and in a watershed treated in 1999 with wollastonite (slow-release Ca source) at the Hubbard Brook Experimental Forest (NH), the nutrition, concentrations of stress related compounds, photosystem function, and sapwood area, were analyzed to assess the influence of Ca addition on the protective biochemistry and carbon (C) capture capacity of foliage during the cold season. Previously reported analysis of the Ca concentrations of a subsample of foliage from the same collections reported here showed higher concentrations in trees in the Ca-addition than in foliage from trees in the reference watershed in both Nov and Feb. Coincident with differences in Ca nutrition, the amino acids alanine (Ala) and γ-aminobutyric acid (GABA), and the polyamines putrescine (Put) and spermidine (Spd) were higher in Nov, and concentrations of Put were higher in Feb, in the foliage of trees from the Ca-addition than concentrations in the foliage of trees from the reference watershed. These polyamines and GABA are thought to have direct protective or other benefits to leaves that may contribute to stress tolerance. The foliage of trees from the Ca-addition had higher total chlorophyll and chlorophyll a concentrations than foliage in trees in the reference watershed in Feb - indicating that Ca-supplemented trees likely retained a greater capacity for winter photosynthesis. In contrast, foliage in trees in the reference had significantly lower glutamic acid (Glu) and higher Ala concentrations than foliage from trees in the Ca-addition watershed in Feb. Similar imbalances in Ala:Glu have been attributed to cold sensitivity or damage in other species. Tree from the Ca-addition watershed also had higher estimated levels of overall foliar biomass than trees from the reference watershed. Together, our findings indicate that Ca-addition resulted in more and better-protected red spruce foliage that had an enhanced capacity for winter C capture compared to the foliage of trees that were subject to ambient soil Ca concentrations. We assessed the causes for year-to-year variations in red spruce foliar winter injury and the influence of injury on xylem growth and carbon (C) sequestration. We used a long-term record of winter injury in a plantation in NH. 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. There was a linear relationship between winter injury and growth reductions for two years after a severe winter injury event. Analysis using data from three New England states indicated that plantation data reflected a regional response. Using regional data we estimated a reduction of 394,000 metric tons of C sequestered in red spruce stems ≥ 20 cm in diameter growing in NY and northern New England during the two years following a severe winter injury event. Reductions in C sequestration likely alter nutrient and water cycling, and other ecosystem services that are influenced by C uptake. PARTICIPANTS: Dr. Paul G. Schaberg and his research team at the Burlington Laboratory of the U.S. Forest Service were major participants in all aspects of this research. We have also worked closely with Dr. C. Eagar of the US Forest Service in Durham, NH as well as numerous researchers (from Harvard, Cornell, Institute of Ecosystem Studies, Boston College) working at Hubbard Brook Experimental Forest in Thornton, NH. There were and continue to be numerous opportunities for training and professional development for several undergraduate students, Masters students and PhD students conducting research associated with the project. TARGET AUDIENCES: Land mangers in the northeastern United States and Canada will benefit from knowledge pertaining to the implications of continued pollution inputs into forested ecosystems that result in nutrient depletions. In addition, policy makers will also benefit from data summarizing the implications of acid-rain induced calcium depletion on our northern forests. Other scientists will also utilize our results to aid them in understanding the implications of calcium depletion including modelers that use such data to predict growth and product losses associated with anthropogenic inputs. PROJECT MODIFICATIONS: Not relevant to this project.

Impacts
This work will provide the first quantitative evaluation of how pollution-influenced winter injury affects the health and productivity of red spruce, and will enable us to quantitatively evaluate the links between Ca nutrition, foliar injury, and changes in forest productivity. This work will provide contextual information for policy makers and managers seeking to understand the potential landscape-scale impacts of continued acidic deposition and Ca depletion, and will provide data needed for possible future work to evaluate the economic impacts of pollution-exacerbated winter injury. This work (combined with past findings) will help evaluate the cause(s) for the decline of an economically and ecologically important tree species, and help assess the possibility that Ca depletion more broadly predisposes trees to inadequate stress response and decline - a scenario with significant management and policy implications for the Northern Forest

Publications

  • Schaberg, P.G.; G.J. Hawley. 2010. Disruption of calcium nutrition at Hubbard Brook Experimental Forest alters the health and productivity of red spruce and sugar maple trees and provides lessons pertinent to other sites and regions. Proceedings of the Conference on the Ecology and Management of High-Elevation Forests of the Central and Southern Appalachian Mountains. May 14-15, Snowshoe Mountain resort, Slatyfork, West Virginia. USDA Forest Service General Technical Report. In press.
  • Schaberg, P.G., Miller, E.K., Eagar, C. 2010. Assessing the Threat that Anthropogenic Calcium Depletion Poses to Forest Health and Productivity. USDA Forest Service General Technical Report and the web-based forestry encyclopedia: www.threats.forestencyclopedia.net. In press.
  • Strimbeck, G.R., Kjellsen, T.D., Schaberg, P.G., Murakami, P.F. 2008. Dynamics of low-temperature acclimation in temperate and boreal conifer foliage in a mild winter climate. Tree Physiol. 28:1365-1374.


Progress 10/01/07 to 09/30/08

Outputs
OUTPUTS: In one study field collections of xylem cores (for growth assessment) and foliage (for nutrient analysis) have been completed. Xylem cores have been fully processed and foliar nutrition measurements are complete and are summarized and combined into a master spreadsheet that for statistical analyses of the influence of winter injury and calcium nutrition on growth. this data is now being used in stepwise linear regression analyses to test if specific weather patterns or events can help explain growth trends found from xylem core analysis - especially growth reductions associated with years of known high winter injury. Once regression analyses are complete, then tables and figures summarizing study results will be constructed and revised, and a manuscript summarizing study findings will be drafted. Once finalized, the draft manuscript will be sent out to four outside reviewers, revised and then submitted to a peer-reviewed journal for possible publication. Study results will also be presented at scientific meetings probably in 2010.In another study field collections of foliage from dominant and co-dominant sugar maple trees were collected at Hubbard Brook in August 2007 and were processed to assess cation nutrition differences associated with treatment. Foliage was also used to measure antioxidant enzyme activity, sugar and starch concentrations and woody shoots associated with foliar harvests were also analyzed for sugar and starch concentrations. Current-year shoots were collected in winter 2008 to measure to cold tolerance and carbohydrate (sugar and starch) concentrations. Data indicates that Ca and Al treatments had significant influences on leaf nutrition and foliar antioxidant enzyme activity. Surprisingly, analyses of shoot tissues showed no treatment-associated differences in cold tolerance, but did show significant differences in carbohydrate levels - notably sugars. Also surprisingly, most treatment-associated differences in leaf and shoot physiology are associated with the Al treatment, suggesting that trees in this treatment are now experiencing meaningful stress. Based on these findings, we added measurements of soil-based extracellular enzyme activity (EEA) in spring and fall to determine if the treatment differences found for trees in summer were accompanied by broader soil-based impacts outside of the standard growing season. Results of these analyses showed that Ca and Al treatment both influenced soil EEA measures, but that the nature and degree of influence varied with the season. Data analysis to date indicate that Al additions are significantly altering tree- and soil-based physiology, and that these changes have occurred at foliar Al concentrations within the range considered normal for healthy sugar maple trees. Results are being written up as the Thesis for a Masters of Science student at UVM to be defended in May 2009. The central core or this Thesis is being written in a journal format, and will be sent to four reviewers in the spring of 2009 prior to submission to a peer-reviewed journal in fall 2009. Results of this study will also be presented at scientific conferences during summer 2009. PARTICIPANTS: Dr. Paul G. Schaberg and his research team at the Burlington Laboratory of the U.S. Forest Service were major participants in all aspects of this research. We have also worked closely with Dr. C. Eagar of the US Forest Service in Durham, NH as well as numerous researchers (from Harvard, Cornell, Institute of Ecosystem Studies, Boston College) working at Hubbard Brook Experimental Forest in Thornton, NH. There were and continue to be numerous opportunities for training and professional development for several undergraduate students, Masters students and PhD students conducting research associated with the project. TARGET AUDIENCES: Land mangers in the northeastern United States and Canada will benefit from knowledge pertaining to the implications of continued pollution inputs into forested ecosystems that result in nutrient depletions. In addition, policy makers will also benefit from data summarizing the implications of acid-rain induced calcium depletion on our northern forests. Other scientists will also utilize our results to aid them in understanding the implications of calcium depletion including modelers that use such data to predict growth and product losses associated with anthropogenic inputs. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
For the first study described above the proposed work will provide the first quantitative evaluation of how pollution-influenced winter injury affects the health and productivity of red spruce, and will enable us to quantitatively evaluate the links between Ca nutrition, foliar injury, and changes in forest productivity. This work will provide contextual information for policy makers and managers seeking to understand the potential landscape-scale impacts of continued acidic deposition and Ca depletion, and will provide data needed for possible future work to evaluate the economic impacts of pollution-exacerbated winter injury. For the second study described above, the research (combined with past findings) will help evaluate the cause(s) for the decline of an economically and ecologically important tree species, and help assess the possibility that Ca depletion more broadly predisposes trees to inadequate stress response and decline - a scenario with significant management and policy implications for the Northern Forest

Publications

  • Huggett, B.A., Schaberg, P.G., Hawley, G.J., Eagar, C. 2007. Long-term calcium addition increases growth release, wound closure and health of sugar maple (Acer saccharum) trees at the Hubbard Brook Experimental Forest. Canadian Journal of Forest Research. 37:1692-1700.
  • Halman, J. M., P.G. Schaberg, G.J. Hawley, C. Eagar. 2008. Calcium addition at the Hubbard Brook Experimental Forest increases sugar storage, antioxidant activity, and cold tolerance in native red spruce (Picea rubens Sarg.). Tree Physiology. 28:855-862.
  • Schaberg, P.G. 2008. Tums for trees. Northern Woodlands. 15:57.