Source: NORTH CAROLINA STATE UNIV submitted to
ECOLOGICAL & ECONOMIC INTEGRITY OF SOUTHERN FOREST ECOSYSTEMS- STAND DYNAMICS, INVASIVE PESTS & GENETIC DETERMINANTS OF KEYSTONE TREES
Sponsoring Institution
National Institute of Food and Agriculture
Project Status
TERMINATED
Funding Source
Reporting Frequency
Annual
Accession No.
0202068
Grant No.
(N/A)
Project No.
NCZ04202
Proposal No.
(N/A)
Multistate No.
(N/A)
Program Code
(N/A)
Project Start Date
Oct 1, 2004
Project End Date
Sep 30, 2010
Grant Year
(N/A)
Project Director
Robison, D. J.
Recipient Organization
NORTH CAROLINA STATE UNIV
(N/A)
RALEIGH,NC 27695
Performing Department
Forestry and Environmental Resources
Non Technical Summary
The Southern Appalachian forests are integral to the health and vitality of a large eastern region. Several keystone tree species, with important ecological and economic functions are threatened by changing conditions and exotic pests. We will study northern red oak, eastern and Carolina hemlock, and Fraser fir. This will investigate ways to promote the regenation of northern red oak, to conserve the germplasm of hemlock, and to understand the genetic and ecological factors involved in Fraser fir pest resistance.
Animal Health Component
(N/A)
Research Effort Categories
Basic
10%
Applied
70%
Developmental
20%
Classification

Knowledge Area (KA)Subject of Investigation (SOI)Field of Science (FOS)Percent
1230621107034%
2110611107056%
2120611107010%
Goals / Objectives
This work will investigate and search for solutions to vexing challenges in Southern Appalachian forested ecosystems, which threaten the integrity, sustainability and vitality of keystone tree species in these landscapes. We will - 1. Determine new ways to promote the growth of the ecologically critical and difficult to regenerate but highly valued northern red oak in mixed species forests through enrichment planting approaches. 2. Examine and conserve the genetic resources of the ecologically critical and threatened eastern hemlock in mixed species forests through genotype sampling and ex-situ conservation of germplasm. 3. Explore the genetics and physiology of the ecologically critical and threatened Fraser fir in high elevation mixed species forests through population studies of Fraser fir and it's balsam wooly adelgid pest. Explore breeding and cultural approaches to controlling Phytophthora root rot in Christmas tree plantings, to sustain and support economic development of rural Appalachian communities.
Project Methods
The southern Appalachian forests run north to south from Virginia to Alabama. They are home to critically important populations of diverse plants and animals, tourism and forest products industries, and represent an ecologically, economically and culturally critical component of the eastern U.S. landscape. These upland forests have been widely mismanaged over several hundred years, particularly up to the early 1900s. For the past 100 years or so they have proved amazing resilient and have recovered from the mistreatment of past agricultural and forest practices (MacCleery 2002). However, with increasingly alarming regularity, these landscapes and the people they serve have suffered convulsions from exotic invasive pests and management limitations which now threaten the stability of these landscapes and the hard-won recovery they have had. Across much of the region there are several keystone tree species which together provide a good deal of the thread which holds the ecosystem stable. While it can be argued which species are most important, three that are surely critical are northern red oak, eastern hemlock and Fraser fir. Each of these is currently challenged. Without focused concerted effort to secure the future of these species the southern Appalachian forests will suffer unpredictable and degrading changes.

Progress 10/01/04 to 09/30/10

Outputs
OUTPUTS: 1. Worked on new ways to promote the growth of the ecologically critical and difficult to regenerate but highly valued northern red oak in mixed species forests through enrichment planting approaches, and better understanding of how oaks in mixed speciaes natural stands interact with a range of other species competing for the same resources. 2. Have continued work on the genetics of the criticaly important and threatened eastern hemlock in mixed species forests throughout the southern Appalachians through genotype sampling and ex-situ conservation of germplasm. Seed has been collected from many stands, tested and germinated, and conservation banks planted to preserve the germplasm. Worked on genetic resource conservation for the critically endangered eastern and Carolina hemlock, ecologically important species in the mixed forests of the eastern and upper-midwestern regions of the United States that are threatened by the invasive hemlock woolly adelgid. 3. Have exmained the genetics and physiology of the ecologically critical and threatened Fraser fir in high elevation mixed species forests through population studies of Fraser fir and it's balsam wooly adelgid pest, and investigated breeding and cultural approaches to controlling Phytophthora root rot in Christmas tree plantings of this same species, to sustain and support economic development of rural Appalachian communities. Refined techniques for Christmas tree growers to graft Fraser fir onto Phytophthora resistance rootstocks. Developed techniques to screen fir species for resistance to BWA, completed microsatellite study of genetic population structure in Fraser and intermediate firs, surveyed the effect of HWA on water relations in hemlocks, ramped up interspecific hybridization of Fraser fir and other firs with adelgid and/or Phytophthora resistance, and did a transcriptome analysis of Fraser fir. PARTICIPANTS: Nothing significant to report during this reporting period. TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
Oak regeneration in mixed hardwood upland stands can be highly variable and difficult to manipulate through silvicultural practices designed to promote its growth and development in the context of natural competition. Enrichment planting studies have indicated that minimal weed control and fertilization are generally insufficient to promote oak growth, but in other studies oaks have responded positively when treated within natural stands from natural regeneration. Growth of oaks and related species has been captured in models that can predict growth under a variety of conditions. Seed collected from hemlocks in the southern Appalahians where it is very much threatened by the hemlock wooley adelgid (HWA), has been successfully germinated, gorwn in nursery conditions to seedling size and planted and maintained in ex-situ conservation banks in South America, and one in the central U.S., both outside of the range of HWA. These plantings now represent a key component of conserving this species while means to manage HWA are pursued. Seeds collected from natural stands of eastern and Carolina hemlock have been germinated and grown in nurseries to the seedling stage and subsequently outplanted into ex situ conservation plantings in Chile, Brazil, and North Carolina (USA). Seedlings of both species are currently being grown in a forest nursery in Arkansas (USA) for eventual conservation bank planting in the Ozark Mountains. Seed collections from natural populations of eastern hemlock have been expanded beyond the southeastern US and conserved genetic resources now include germplasm from significant portions of the northeastern and upper-midwestern regions of the US. Genetic structure and diversity studies have been completed for both hemlock species and results are being utilized to refine continued seed collection strategies and management of planted ex situ conservation banks. The genetics and physiology of the ecologically critical and threatened Fraser fir in high elevation mixed species forests have been studied and it and it's pest balsam wooley adelgid (BWA) can now be characterized by discrete and related population structures. This information provides the basis for understanding the origins of this pest, how it's variation interacts with hosts (Fraser fir) genetic variation, and clues as to where and how genetically based pest and tree interaction management might be developed. Phytophthora root rot in Fraser fir in Christmas tree plantings have also been studied, in order to understand this alternative and important pest of Fraser fir both for ecological and economic reasons.

Publications

  • Williams, JL. 2009. Predatory Mites (Acari: Phytoseiidae) in Fraser fir Christmas Tree Plantations in the Southern Appalachians of North Carolina. PhD Thesis. NC State University, Raleigh, NC. 116 pp.
  • Kim, YW, R Newton, J Frampton and KH Han. 2009. Embryogenic tissue initiation and somatic embryogenesis in Fraser fir (Abies fraseir [Pursh] Poir.). In Vitro Cellular and Developmental Biology - Plant. 45:400-406.
  • Schuler, JL and DJ Robison. 2008. The effects of intensive management on the leaf characteristics and growth phenology of young yellow-poplar stems. For Ecol Manage. 255:787-796. McTague, JP, D O'Loughlin, JP Roise, DJ Robison, RC Kellison. 2008. The SOHARC Model System for Growth and Yield of Southern Hardwoods. So J of App For. 32:173-183.
  • Potter, KM, WS Dvorak, BS Crane, VD Hipkins, RM Jetton, WA Whittier and R Rhea. 2008. Allozyme variation and recent evolutionary history of eastern hemlock (Tsuga canadensis) in the southeastern United States. New For. 35:131-145.
  • RM Jetton, WS Dvorak and WA Whittier. 2008. Ecological and genetic factors that define the natural distribution of Carolina hemlock in the southeastern United States and their role in ex situ conservation. For. Ecol. Manage. 255: 3212-3221
  • Jetton, RM, WA Whittier, WS Dvorak and KM Potter. 2008. StAtuS oF ex situ conServAtion eFFortS For cArolinA And eAStern hemlock in the SoutheAStern united StAteS. Fourth symposium on Hemlock Woolly adelgid in the Eastern United states. USDA Forest Service Notes: 81-88.
  • Jetton, RM. 2008. Biological Control, Host Resistance, and Vegetative Propagation: Strategies and Tools for Management of the Invasive Hemlock Woolly Adelgid, Adelges tsugae Annand. PhD Thesis, North Carolina State University, Raleigh, NC. 211 pp.
  • Potter, KM, J Frampton, SA Josserand, DC Nelson, C. D. 2008. Genetic variation and population structure in Fraser fir (Abies fraseri): a microsatellite assessment of young trees. Can J For Res. 38:2128-2137.
  • Gocke, M.H., D.J. Robison and B. Goldfarb. 2006. The effect of prune location on new shoot growth and subsequent rooting ability for containerized northern red oak seedling stock plants. So. J. Appl. For. Submitted.
  • Schuler, J.L., D.J. Robison, R. Myers, and M.J. Young. 2006. Reproduction Responses of Southern Hardwood Forests to Full and Partial Overstory Removals. American Midland Naturalist. Accepted pending revision.
  • Schuler, J.L. and D.J. Robison. 2006. Stand Development and Growth Response of 1 to 3 Year-Old Natural Upland Hardwoods to Silvicultural Treatments. Forest Ecology and Manage. 232:124-134.
  • Stejskal, J. J. Kobliha, and J. Frampton. 2011. Results of Czech-American cooperation in interspecific fir hybridizaiton in 2008 and 2009. Czech J. For. Sci. 57(3):114-122.
  • Jetton, R.M., W.A. Whittier, W.S. Dvorak, and J.R. Rhea. 2010. Status of gene conservation for eastern and Carolina hemlock in the eastern United States. pp. 93-99 In: B. Onken and R. Reardon (Eds.), Proceedings of the Fifth Symposium on Hemlock Woolly Adelgid in the Eastern United States.
  • Hibbert-Frey, H., J. Frampton, F. Blazich, D. Hundley, and E. Hinesley. 2011. Grafting Fraser fir: effect of scion origin (crown position and branch order). HortScience. 46(1):91-94.
  • Newton, L., J. Frampton, J. Monahan, B. Goldfarb, F. Hain. 2011. Two novel techniques to screen Abies seedlings for resistance to the balsam woolly adelgid, Adelges piceae. J. of Insect Sci. 11.158 available online: insectscience.org/11.158.
  • Whetten, R., J. Frampton, and N. Gaddis. 2011 Transcriptome analysis of Fraser fir.10th International Christmas Tree Research & Extension Conf. Eichgraben, Austria, August 20 ‐ 26. (abstract and presentation)
  • Rhea, J.R. and R. Jetton. 2010. Framework for genetic conservation of eastern (Tsuga canadensis) and Carolina (T. caroliniana) hemlock. pp. 106-121 In: B. Onken and R. Reardon (Eds.), Proceedings of the Fifth Symposium on Hemlock Woolly Adelgid in the Eastern United States. Potter, K.M., R.M.
  • Jetton, W.S. Dvorak, J. Frampton, and J.R. Rhea. 2010. Ex situ seed collection represents genetic variation present in natural stands of Carolina hemlock. pp. 181-190 In: B. Onken and R. Reardon (Eds.), Proceedings of the Fifth Symposium on Hemlock Woolly Adelgid in the Eastern United States.
  • Schuler, J and D Robison. 2010. Performance of northern red oak enrichment plantings in naturally regenerating southern Appalachian hardwood stands. New Forests 40:119-130.
  • Hibbert-Frey, H., J. Frampton, F.A. Balzich and L.E. Hinesley. 2010. Grafting Fraser fir (Abies fraseri): Effect of grafting date, shade, and irrigation. HortScience. 45(4):617-620.
  • Potter, K.M., J. Frampton, S.A. Josserand and C.D. Nelson. 2010. Evolutionary history of two imperiled Southern Appalachian conifers, Fraser fir and intermediate fir, revealed using microsatellite markers. Conservation Genetics. 11:1499-1513.
  • Potter, K.M., R.M. Jetton, W.S. Dvorak, J. Frampton and R. Rhea. 2010. Ex situ seed collection represents genetic variation present in natural stands of Carolina hemlock. p181-190, in: Onken, B. and R. Reardon, eds., Fifth Symp. on Hemlock Woolly Adelgid in the Eastern United States. USDA For. Serv. For. Health Protection. Morgantown, WV. 222p
  • Rivera L.N, J.-C. Domec, J. Frampton, F. Hain, I. Peszlen, and J.S. King. 2010. The effect of hemlock woolly adelgid (HWA) infestation on water relations of Carolina and eastern hemlock: Can ecophysiological investigation of tree water relations improve silvicultural management of the HWA p129-131, in: Onken, B. and R. Reardon, eds., Fifth Symp. on Hemlock Woolly Adelgid in the Eastern United States. USDA For. Serv. For. Health Protection. Morgantown, WV. 222p
  • Berenguer, BJ, JL Schuler, DJ Robison and MH Gocke. 2009. Response of a rising 2 year old natural Piedmont upland stand to fertilization. New Forests 37:155-174.
  • Jetton, R, W Dvorak, A Whittier, K Potter, and R Rhea. 2009. Genetics And Conservation Of Hemlock Species Threatened By The Hemlock Woolly Adelgid. USDA Research Forum on Invasive Species. USDA Forest Service Notes: 39-40
  • Ross W. Whetten, Valenzuela A Sofia, and John Frampton. 2009. Polymerase Chain Reaction Preparation of Template for Massively Parallel Pyrosequencing. J Biomol Tech. 20: 128-134.
  • Walker-Lane, LN. 2009. The Effect of Hemlock Woolly Adelgid Infestation on Water Relations of Carolina and Eastern Hemlock. MS Thesis. NC State University, Raleigh, NC. 61 pp.
  • Frey, HH. 2009. Factors affecting graft success and early growth of Fraser fir. MS Thesis. NC State University, Raleigh, NC. 56 pp.
  • Rutledge, ME, J Frampton, LE Hinesley. 2008. Using a bud volume index with the top-stop nipper to control leader growth of Fraser Fir Christmas trees. HortTechnology.
  • Kaur, N. 2008. Developing Artificial Rearing Techniques for Hemlock Woolly Adelgid, Adelges tsugae and Balsam Woolly Adelgid, Adelges piceae; Artificial Infestation and Epicuticular Wax Study of Carolina Hemlock, Tsuga caroliniana, Provenances. MS Thesis. NC State University, Raleigh, NC. 118 pp.
  • Emerson J L , J Frampton J , SE McKeand. 2008. Genetic Variation in Early Growth and Bud Production among Natural Populations of Fraser Fir. HortScience 43: 661-666.
  • Rutledge, ME, J Frampton, LE Hinesley. 2008. Top-Stop Nipper Reduces Leader Growth in Fraser Fir Christmas Trees. HortTechnology.
  • Aspinwall, ME. 2008. Chemical and Mechanical Methods to Reduce Leader Growth in Fraser Fir. MS Thesis. NC State University, Raleigh, NC. 81 pp.
  • Berenguer, BJ, MH Gocke, JL Schuler, E Treasure and DJ Robison. 2008. Accelerating development with fertilization in a young natural piedmont mixed hardwood-pine stand. In: Proc. 14th Biennial Southern Silvicultural Research Conference. Feb. 2007, Athens, GA. In Press.
  • Potter, K.M., W.S. Dvorak, B.S. Crane, V.D. Hipkins, R. M. Jetton, W.A. Whittier, and R. Rhea. 2008. Allozyme variation and recent evolutionary history of eastern hemlock (Tsuga canadensis) in the southeastern United States. New Forests. (In press)
  • Jetton, R.M., W.S. Dvorak, and W.A. Whittier. 2008. Ecological and Genetic Factors that Define the Natural Distribution and Carolina Hemlock in the Southeastern United States and Their Role in Ex Situ Conservation. Forest Ecology and Management (in press).
  • Balakshin M., E. Capanema, S. Kelley, R. Whetten and J. Frampton. 2008. A methodological approach to elucidate the effect of balsam woolly adelgid infestation on the structure of wood and bark. Conference of the Partnership for Saving Threatened Forests. N.C. State University. Raleigh. (presentation and abstract)
  • Berenguer, BJ, JL Schuler; DJ Robison, MH Gocke and E Treasure. 2008. Response of a very young natural piedmont upland stand to fertility. Accepted pending revision: New Forests.
  • Schuler, JL and DJ Robison. 2008. The Effects of Intensive Management on the Leaf Characteristics and Growth Phenology of Young Yellow-Poplar Stems. Forest Ecology and Manage. 255:787-796.
  • Gocke, MH, H Dreps, E Treasure and DJ Robison. 2008. Maximizing northern red oak rooted cutting opportunities. In: Proc. 14th Biennial Southern Silvicultural Research Conference. Feb. 2007, Athens, GA. In Press.
  • Emerson J., L. van Zyl, J. Frampton and R. Whetten. 2008. Fraser Fir and the Balsam Woolly Adelgid: Applying Microarray Technology to Understand Host-Pest Resistance Interactions. Conference of the Partnership for Saving Threatened Forests. N.C. State University. Raleigh. (presentation and abstract)
  • Frampton J. 2008. Hemlock Response to HWA Infestation. Conference of the Partnership for Saving Threatened Forests. N.C. State University. Raleigh. (presentation and abstract)
  • Whetten, R. and J. Frampton. 2008. High throughput DNA sequencing of Fraser fir (Abies fraseri) p14-15 in: Thomsen, I.M., Rasmussen, H.N. & Sorensen, J.M. (Eds.), Proceedings of the 8th International Christmas Tree Research and Extension Conference. Forest & Landscape Working Papers No. 26-2008. Forest & Landscape Denmark, Horsholm. 138p. (poster presentation and abstract)
  • McTague, J.P., D. OLoughlin, J.R. Roise, D.J. Robison and R.C. Kellison. 2008. The NC State SOHARC model system for growth and yield of Southern hardwoods. So. J. Appl. For.
  • Camcore Annual Report 2006. Genetic Diversity of Carolina and Eastern hemlock. North Carolina State University. pp. 16
  • The Fight to Save the Easts Hemlocks. 2006. Blue Ridge Country Magazine. September/October 2006
  • NC State Tree Preservation Program Working to Save Hemlocks from Extinction. 2006. NC State News Services. September 7, 2006
  • Scientists Try to Save Hemlocks. 2006. News and Observer, Raleigh, NC October 29, 2006.


Progress 01/01/09 to 12/31/09

Outputs
OUTPUTS: 1. Work has continued on the finding new ways to promote the growth of the ecologically critical and difficult to regenerate but highly valued northern red oak in mixed species forests through enrichment planting approaches, and better understanding of how oaks in mixed speciaes natural stands interact with a range of other species competing for the same resources. 2. Have continued work on the genetocs of the criticaly important and and threatened eastern hemlock in mixed species forests thorughout the southern Appalachians through genotype sampling and ex-situ conservation of germplasm. Seed has been collected from many stands, tested and germinated, and conservation banks planted to preserve the germplasm. 3. Have exmained the genetics and physiology of the ecologically critical and threatened Fraser fir in high elevation mixed species forests through population studies of Fraser fir and it's balsam wooly adelgid pest, and investigated breeding and cultural approaches to controlling Phytophthora root rot in Christmas tree plantings of this same species, to sustain and support economic development of rural Appalachian communities. PARTICIPANTS: Nothing significant to report during this reporting period. TARGET AUDIENCES: Information generated by this work has reached many forestry scientists through publications and presentations, and users groups in the southern Appalachians intersted in Fraser fir and hemlocks as species threatened by the exotic invasive BWA and HWA pests. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
1. Oak regeneration has been studied in a number of mixed hardwood upland stands and found to be highly variable and difficult to manipulate through silvicultural practices designed to promote its growth and development in the context of natural competition. Enrichment planting studies have indicated that minimal weed control and fertilization are generally insufficient to promote oak growth, but in otehr studies oaks have responded positively when treated within natural stands from natural regeneration. Growth of oaks and related species has been captured in models that can predict growth under a variety of conditions. Seed collected from hemlocks in the souther Appalahians where it is very much threatened by the hemlock wooley adelgid (HWA), has been successfully germinated, gorwn in nursery conditions to seedling size and planted and maintained in ex-situ conservation banks in South America, and one in the central U.S., both outside of the range of HWA. These plantings now represent a key component of conserving this species while means to manage HWA are pursued. The genetics and physiology of the ecologically critical and threatened Fraser fir in high elevation mixed species forests have been studied and it and it's pest balsam wooley adelgid (BWA) can now be characterized by discrete and related population structures. This information provides the basis for understanding the origins of this pest, how it's variation interacts with hosts (Fraser fir) genetic variation, and clues as to where and how genetically based pest and tree interaction management might be developed. Phytophthora root rot in Fraser fir in Christmas tree plantings have aslo been studied, in order to understand this alternative and important pest of Fraser fir both for ecological and economic reasons.

Publications

  • Berenguer, BJ, JL Schuler, DJ Robison, MH Gocke and E Treasure. 2009. Response of a rising 2-year-old natural Piedmont upland stand to fertilization. New For. 37:155-174.
  • Jetton, R, W Dvorak, A Whittier, K Potter, and R Rhea. 2009. Genetics And Conservation Of Hemlock Species Threatened By The Hemlock Woolly Adelgid. USDA Research Forum on Invasive Species. USDA Forest Service Notes: 39-40
  • Ross W. Whetten, Valenzuela A Sofia, and John Frampton. 2009. Polymerase Chain Reaction Preparation of Template for Massively Parallel Pyrosequencing. J Biomol Tech. 20: 128-134.
  • Walker-Lane, LN. 2009. The Effect of Hemlock Woolly Adelgid Infestation on Water Relations of Carolina and Eastern Hemlock. MS Thesis. NC State University, Raleigh, NC. 61 pp.
  • Frey, HH. 2009. Factors affecting graft success and early growth of Fraser fir. MS Thesis. NC State University, Raleigh, NC. 56 pp.
  • Williams, JL. 2009. Predatory Mites (Acari: Phytoseiidae) in Fraser fir Christmas Tree Plantations in the Southern Appalachians of North Carolina. PhD Thesis. NC State University, Raleigh, NC. 116 pp.
  • Kim, YW, R Newton, J Frampton and KH Han. 2009. Embryogenic tissue initiation and somatic embryogenesis in Fraser fir (Abies fraseir [Pursh] Poir.). In Vitro Cellular and Developmental Biology - Plant. 45:400-406.


Progress 01/01/08 to 12/31/08

Outputs
OUTPUTS: Hemlock Seed Collections was focused on Eastern hemlock, although explorations for additional populations of Carolina hemlock continued. Range-wide site explorations and seed collections have been conducted. Among Eastern hemlock populations were outlier populations in AL and KY that may be of unique genetic interest. Due to drought and the advancing HWA infestation, hemlock seed production has been highly variable across the southeast. Cone-bearing trees have been found in populations located in GA, TN, VA, NC and SC. Seed was collected from mother trees of Carolina hemlock and Eastern hemlock. Scion from surviving mature Fraser fir has been grafted to make material available for testing the degree of genetic resistance to balsam woolly adelgid (BWA). Preliminary tests for genetic variation in resistance of Turkish fir to Phytophthora cinnamomi (Pc) have been completed. Seedlings representing over 100 O-P families of Abies bornmuelleriana and equi-trojani are being cultured. These seedlings are being inoculated with Pc to look at within species variation for resistance and to identify resistant candidates for research. Analysis of gene expression changes in Fraser fir in response to BWA or Pc requires background information on the genes expressed in Fraser fir during normal growth and during disease. To gain information on Fraser fir genes, samples of fir cDNAs have been analyzed by high-throughput sequencing, providing a collection of Fraser fir gene sequences that will underpin efforts to understand genetic responses of fir to a variety of biotic and abiotic stresses, and putative genetic variation to characterize Fraser fir populations. Experiments to measure differences in gene expression due to BWA infestation of Fraser fir are in progress. A variety of objectives are being pursued with regard to oaks and our abilitiy to understand their ecology and to manage them for reproduction, growth and representation in forests. We have continued work to develop growth and yield models for predicting oak growth, experiments have been deployed to learn about the ecological constraints to tree regeneration and competitive ability, and we have pursued vegetatative propagation strategies for oak and enrichment planting approaches so that elite oak genotypes can be planted into natural forests in order to enhance oak representation in places where oak regeneration has been low. PARTICIPANTS: Nothing significant to report during this reporting period. TARGET AUDIENCES: The target audiences for this work are landowners and forestry professionals in the regions where the species being studies are found, and forestry scientists state-, nation-, and world-wide who are interested in these topics. Specifically this information has been provided through outreach efforts to Christmas tree growers and land managers (public and private) in western NC, and provided to many through publications. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
This work collectively provides insight into key components of the souuthern Appalachian forests. The work on Fraser fir and the hemlocks is critical to developing conservation and proactive management plans for species threatened by exotic pests. This work will also provide clues for the management of other ecosystem components similarly threatened by exotic invasive pests. The work with oaks and colocated species addresses a key concern of ecologists and foresters in this region - namely the dificulty in regenerating oak to grow rapidly as a timber and mast producing species of great interest and beauty. As this work progresses it will enhance our capacity to predict forest development and manage to advantage. Ex Situ Conservation of Hemlocks Using seed collected in previous years, work has begun on the establishment of ex-situ conservation banks of hemlock on the lands of Forestal Arauco and CMPC Forestal in Chile and Rigesa in Brazil. All of these companies are long-term members of the Camcore international gene conservation program at NC State. Seedlings are being grown in greenhouses, and field establishment of conservation banks is ongoing. We have finalized growth and yield equations for oak and other species in mixed forests and can now track and predict the growth of those species from simple inventory data. We have begun to understand the contributions of spacing, fertility and other factors on the regeneration and success of inidividual oaks and other species, and population level resposnes as well. A rooted cutting system has been developed so that elite oak genotypes can be cloned and an enrichment planting system is under development to deploy these types of trees into otherwise naturally regenerated stands. We anticipate that other sceintists and field managers will benefit from this emerging work and be better able to manage our natural forests for this critical species.

Publications

  • Schuler, JL and DJ Robison. 2008. The effects of intensive management on the leaf characteristics and growth phenology of young yellow-poplar stems. For Ecol Manage. 255:787-796. McTague, JP, D O'Loughlin, JP Roise, DJ Robison, RC Kellison. 2008. The SOHARC Model System for Growth and Yield of Southern Hardwoods. So J of App For. 32:173-183.
  • Potter, KM, WS Dvorak, BS Crane, VD Hipkins, RM Jetton, WA Whittier and R Rhea. 2008. Allozyme variation and recent evolutionary history of eastern hemlock (Tsuga canadensis) in the southeastern United States. New For. 35:131-145.
  • RM Jetton, WS Dvorak and WA Whittier. 2008. Ecological and genetic factors that define the natural distribution of Carolina hemlock in the southeastern United States and their role in ex situ conservation. For. Ecol. Manage. 255: 3212-3221
  • Jetton, RM, WA Whittier, WS Dvorak and KM Potter. 2008. StAtuS oF ex situ conServAtion eFFortS For cArolinA And eAStern hemlock in the SoutheAStern united StAteS. Fourth symposium on Hemlock Woolly adelgid in the Eastern United states. USDA Forest Service Notes: 81-88.
  • Jetton, RM. 2008. Biological Control, Host Resistance, and Vegetative Propagation: Strategies and Tools for Management of the Invasive Hemlock Woolly Adelgid, Adelges tsugae Annand. PhD Thesis, North Carolina State University, Raleigh, NC. 211 pp.
  • Potter, KM, J Frampton, SA Josserand, DC Nelson, C. D. 2008. Genetic variation and population structure in Fraser fir (Abies fraseri): a microsatellite assessment of young trees. Can J For Res. 38:2128-2137.
  • Kaur, N. 2008. Developing Artificial Rearing Techniques for Hemlock Woolly Adelgid, Adelges tsugae and Balsam Woolly Adelgid, Adelges piceae; Artificial Infestation and Epicuticular Wax Study of Carolina Hemlock, Tsuga caroliniana, Provenances. MS Thesis. NC State University, Raleigh, NC. 118 pp.
  • Rutledge, ME, J Frampton, LE Hinesley. 2008. Using a bud volume index with the top-stop nipper to control leader growth of Fraser Fir Christmas trees. HortTechnology.
  • Emerson J L , J Frampton J , SE McKeand. 2008. Genetic Variation in Early Growth and Bud Production among Natural Populations of Fraser Fir. HortScience 43: 661-666.
  • Rutledge, ME, J Frampton, LE Hinesley. 2008. Top-Stop Nipper Reduces Leader Growth in Fraser Fir Christmas Trees. HortTechnology.
  • Aspinwall, ME. 2008. Chemical and Mechanical Methods to Reduce Leader Growth in Fraser Fir. MS Thesis. NC State University, Raleigh, NC. 81 pp.


Progress 01/01/07 to 12/31/07

Outputs
OUTPUTS: 2007 Hemlock Seed Collections was focused on Eastern hemlock, although explorations for additional populations of Carolina hemlock continued. Range-wide site explorations were conducted June to Aug 2007 and seed collections in Sept and Oct. Among Eastern hemlock populations were outlier populations in AL and KY that may be of unique genetic interest. Due to drought and the advancing HWA infestation, hemlock seed production was sparse across the southeast in 2007. However, cone-bearing trees were found in populations located in GA, TN, VA, NC and SC. Seed was collected from 6 mother trees in one provenance of Carolina hemlock and 69 mother trees in 14 provenances of Eastern hemlock. Scion from surviving mature Fraser fir has been grafted, but is growing slowly, so available material is not sufficient for testing the degree of genetic resistance to balsam woolly adelgid (BWA). We have developed methods to inoculate fir seedlings and grafts with BWA. A preliminary test for genetic variation in resistance of Turkish fir to Phytophthora cinnamomi (Pc) has been completed. About 3,500 seedlings representing over 100 O-P families of Abies bornmuelleriana and equi-trojani are being cultured in the greenhouse. These seedlings will be inoculated with Pc to look at within species variation for resistance and to identify resistant candidates for research. Analysis of gene expression changes in Fraser fir in response to BWA or Pc requires background information on the genes expressed in Fraser fir during normal growth and during disease. To gain information on Fraser fir genes, 3 samples of fir cDNAs were submitted for high-throughput sequencing. This provided a collection of about 30,000 Fraser fir gene sequences that will underpin efforts to understand genetic responses of fir to a variety of biotic and abiotic stresses, and over 15,000 sites of putative genetic variation to characterize Fraser fir populations. Experiments to measure differences in gene expression due to BWA infestation of Fraser fir are in progress, and analysis of these experiments should be completed in 2008. A variety of objectives are being pursued with regard to oaks and our abilitiy to understand their ecology and to manage them for reproduction, growth and representation in forests. We have continued work to develop growth and yield models for predicting oak growth, experiments have been deployed to learn about the ecological constraints to tree regeneration and competitive ability, and we have pursued vegetatative propagation strategies for oak and enrichment planting approaches so that elite oak genotypes can be planted into natural forests in order to enhance oak representation in places where oak regeneration has been low. PARTICIPANTS: Nothing significant to report during this reporting period. TARGET AUDIENCES: Nothing significant to report during this reporting period. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
This work will collectively provide greater scientific insight into some of the key components of the souuthern Appalachian forests. The work on Fraser fir and the hemlocks is critical to developing conservation and proactive management plans for species threatened by exotic pests. This work will also provide clues for the management of other ecosystem components similarly threatened by exotic invasive pests. The work with oaks addresses a key concern of ecologists and foresters in this region - namely the dificulty in regenerating oak to grow rapidly as a timber and mast producing species of great interest and beauty. As this work progresses it will enhance our capacity to predict forest development and manage to advantage. Ex Situ Conservation of Hemlocks Using seed collected in previous years, work has begun on the establishment of ex-situ conservation banks of hemlock on the lands of Forestal Arauco and CMPC Forestal in Chile and Rigesa in Brazil. All of these companies are long-term members of the Camcore international gene conservation program at NC State. Seedlings are being grown in greenhouses, and field establishment of conservation banks should begin in 2009 and 2010. In addition, the possibility of establishing a conservation bank at the Ouachita National Forest and/or the Ozark National Forest in Arkansas is being investigated. We have finalized growth and yield equations for oak and other species in mixed forests and can now track and predict the growth of those species from simple inventory data. We have begun to understand the contributions of spacing, fertility and other factors on the regeneration and success of inidividual oaks and other species, and population level resposnes as well. A rooted cutting system has been developed so that elite oak genotypes can be cloned and an enrichment planting system is under development to deploy these types of trees into otherwise naturally regenerated stands. We anticipate that other sceintists and field managers will benefit from this emerging work and be better able to manage our natural forests for this critical species.

Publications

  • Schuler, J.L. and D.J. Robison. 2006. Stand Development and Growth Response of 1 to 3 Year-Old Natural Upland Hardwoods to Silvicultural Treatments. Forest Ecology and Manage. 232:124-134.
  • Berenguer, BJ, MH Gocke, JL Schuler, E Treasure and DJ Robison. Accelerating development with fertilization in a young natural piedmont mixed hardwood-pine stand. In: Proc. 14th Biennial Southern Silvicultural Research Conference. Feb. 2007, Athens, GA. In Press.
  • Potter, K.M., W.S. Dvorak, B.S. Crane, V.D. Hipkins, R. M. Jetton, W.A. Whittier, and R. Rhea. 2008. Allozyme variation and recent evolutionary history of eastern hemlock (Tsuga canadensis) in the southeastern United States. New Forests. (In press)
  • Jetton, R.M., W.S. Dvorak, and W.A. Whittier. 2008. Ecological and Genetic Factors that Define the Natural Distribution and Carolina Hemlock in the Southeastern United States and Their Role in Ex Situ Conservation. Forest Ecology and Management (in press).
  • Balakshin M., E. Capanema, S. Kelley, R. Whetten and J. Frampton. 2008. A methodological approach to elucidate the effect of balsam woolly adelgid infestation on the structure of wood and bark. Conference of the Partnership for Saving Threatened Forests. N.C. State University. Raleigh. (presentation and abstract)
  • Berenguer, BJ, JL Schuler; DJ Robison, MH Gocke and E Treasure. 2008. Response of a very young natural piedmont upland stand to fertility. Accepted pending revision: New Forests.
  • Schuler, JL and DJ Robison. 2008. The Effects of Intensive Management on the Leaf Characteristics and Growth Phenology of Young Yellow-Poplar Stems. Forest Ecology and Manage. 255:787-796.
  • Gocke, MH, H Dreps, E Treasure and DJ Robison. 2008. Maximizing northern red oak rooted cutting opportunities. In: Proc. 14th Biennial Southern Silvicultural Research Conference. Feb. 2007, Athens, GA. In Press.
  • Emerson J., L. van Zyl, J. Frampton and R. Whetten. 2008. Fraser Fir and the Balsam Woolly Adelgid: Applying Microarray Technology to Understand Host-Pest Resistance Interactions. Conference of the Partnership for Saving Threatened Forests. N.C. State University. Raleigh. (presentation and abstract)
  • Frampton J. 2008. Hemlock Response to HWA Infestation. Conference of the Partnership for Saving Threatened Forests. N.C. State University. Raleigh. (presentation and abstract)
  • Whetten, R. and J. Frampton. 2008. High throughput DNA sequencing of Fraser fir (Abies fraseri) p14-15 in: Thomsen, I.M., Rasmussen, H.N. & Sorensen, J.M. (Eds.), Proceedings of the 8th International Christmas Tree Research and Extension Conference. Forest & Landscape Working Papers No. 26-2008. Forest & Landscape Denmark, Horsholm. 138p. (poster presentation and abstract)


Progress 01/01/06 to 12/31/06

Outputs
Scion from surviving mature Fraser fir has been grafted, but is growing extremely slowly, so the available materials is not yet sufficient for testing the degree of genetic resistance to balsam woolly adelgid (BWA). A preliminary test for genetic variation in resistance of Turkish fir to Phytophthora cinnamomi (Pc) has been completed, and a second experiment to confirm and extend the results of the preliminary test is underway. Analysis of gene expression changes in Fraser fir in response to either BWA or Pc requires background information on the genes expressed in Fraser fir during normal growth and during a disease response. To gain information on Fraser fir genes, three samples of fir cDNAs have been submitted for high-throughput sequencing. This experiment is still in progress, but if it is successful, it will provide a collection of Fraser fir gene sequences that will underpin future efforts to understand genetic responses of fir to a wide variety of biotic and abiotic stresses. Research is underway on the ecological determinants of oak regeneration and success in naturally regenerated and planted systems. A study of seedling oaks in competition with seedling yellow poplar, in order to evaluate the impacts of belowground competition yielded only preliminary data due to high levels of variation. Another study of the response of natural regeneration to fertilization and thinning has revealed significant means to relieve competition for resources and enhance growth. Explorations for potential Eastern hemlock seed collection sites in June and July revealed that there was very little cone production in the southeastern US in 2006. Still, some seed was collected for the establishment ex situ conservation areas. Soil analysis for both Eastern and Carolina hemlock to better understand the ecology of both species has been accomplished. We have completed a genetic assessment of Eastern hemlock in collaboration with the US Forest Service. Results indicate that populations along the eastern periphery and in the Appalachian Mountains exhibit higher levels of isozyme variation than compared to those along the western periphery of the species range.

Impacts
This work will collectively provide greater scientific insight into some of the key components of the souuthern Appalachian forests. The work on Fraser fir and the hemlocks is critical to developing conservation and proactive management plans for species threatened by exotic pests. This work will also provide clues for the management of other ecosystem components similarly threatened by exotic invasive pests. The work with oaks addresses a key concern of ecologists and foresters in this region - namely the dificulty in regenerating oak to grow rapidly as a timber and mast producing species of great interest and beauty. As this work progresses it will enhance our capacity to predict forest development and manage to advantage.

Publications

  • Camcore Annual Report 2006. Genetic Diversity of Carolina and Eastern hemlock. North Carolina State University. pp. 16
  • The Fight to Save the Easts Hemlocks. Blue Ridge Country Magazine. September/October 2006
  • NC State Tree Preservation Program Working to Save Hemlocks from Extinction. NC State News Services. September 7, 2006
  • Scientists Try to Save Hemlocks. News and Observer, Raleigh, NC October 29, 2006.
  • K.M. Potter, W.S. Dvorak, B.S. Crane, V.D. Hipkins, R.M. Jetton, W.A.Whittier, and R. Rhea. 2007. Genetic Variation and Recent Evolutionary History of Eastern Hemlock (Tsuga canadensis) in the Southeastern United States (submitted to New Forests)
  • Schuler, JL and DJ Robison. Performance of northern red oak enrichment plantings in naturally regenerating Southern Appalachian hardwood stands. In preparation 2007.
  • Berenguer, BJ, DJ Robison, JL Schuler and MH Gocke. Fertilization impacts on growth and species compositiomn in a very young naturally regenerated Peidmont upland hardwood stand in NC. In preparation 2007.
  • Gocke, M.H., D.J. Robison and B. Goldfarb. 2006. The effect of prune location on new shoot growth and subsequent rooting ability for containerized northern red oak seedling stock plants. So. J. Appl. For. Submitted.
  • McTague, J.P., D. OLoughlin, J.R. Roise, D.J. Robison and R.C. Kellison. 2006. The NC State SOHARC model system for growth and yield of Southern hardwoods. So. J. Appl. For. Accepted pending revision.
  • Schuler, J.L., D.J. Robison, R. Myers, and M.J. Young. 2006. Reproduction Responses of Southern Hardwood Forests to Full and Partial Overstory Removals. American Midland Naturalist. Accepted pending revision.