NE1015: Biological Improvement, Habitat Restoration, and Horticultural Development of Chestnut by Management of Populations, Pathogens, and Pests
- Duration:
- October 01, 2003 to September 30, 2008
- Administrative Advisor(s):
-
Michael E. Vayda
(VT.)
- NIFA Reps:
-
Robert M. Nowierski
Statement of Issue(s) and Justification:
Before the introduction of chestnut blight, American chestnut was one of the most valuable trees in eastern North American forests. When Europeans first settled eastern North America, chestnut was a major and eminently notable forest tree everywhere from Ontario to Georgia. It was the one eastern forest tree that approached the dimensions of the gigantic Pacific Coast rain forest conifers; thus, it was referred to as the 'Redwood of the East'. Chestnut occupied a wide range of site types, ranging from mesic to xeric. Quality chestnut on mesic sites attained a clear trunk length of up to 72 feet with an overall height of 120 feet and a diameter of 5 feet. Dry, thin, acidic ridge top soils provided the poorest growing conditions, but chestnut still managed to grow to a height of 70 feet. The tree was prized for its high quality wood, tannin extracts, and nuts. Chestnut blight disease eliminated the American chestnut as a canopy species and elevated oaks as the most dominant tree group in the southern Appalachians. However, widespread oak decline has occurred in the southeastern U.S. over the past 20 years, leading to declining forests. It is believed that southern Appalachian forest ecosystems will not be considered healthy until chestnut can be reintroduced as a functioning component of that ecosystem. Loss of the American chestnut resulted in significant biological and economic changes. Replacement species generally include red oak, black oak, and chestnut oak, all much slower growing than chestnut. Return of chestnut to eastern forests could vastly improve existing forest stands, and positively impact wildlife, since chestnuts produce seed every year as opposed to oaks and hickories.The NE-140 Technical Committee was organized in 1982, and since then, members have worked cooperatively to investigate the complex issues involved with the biological improvement of chestnut, along with habitat restoration and horticultural development of American chestnut and chestnut cultivars. Environmental and land stewardship issues that are concerns to stakeholders are addressed by NE-140 through efforts to understand the ecology and biology of eastern North American forests following the loss of American chestnut. Control of chestnut blight disease through the use of hypoviruses and resistant chestnut trees may vastly improve forest stands in eastern North America, answering many concerns regarding forest ecology. Other important issues of this continued work are improved forest management, including biological-based controls of invasive insect and disease organisms threatening chestnut. This will serve as a model for natural resource managers. Food safety issues are addressed via the creation and use of IPM-based controls of both forest and orchard trees, reducing the use of pesticides. Increased use of chestnuts in the American diet will improve human health, since chestnuts are unique among nuts with fat content ranges from 1-10%, compared to 70% in most other nuts. The viability of rural communities will be enhanced through creation of a new horticultural chestnut market.
Restoration of chestnut as a timber tree and improving orchard chestnuts in the United States continue to be the long-term goals of the members of this regional project. To succeed in our efforts to achieve these long-term goals, continued activity of the award-winning NE-140 group is critical. NE-140 is working to successfully develop and deploy biological systems that protect natural resources and the environment. NE-140 must sustain its rapid, productive trajectory in order to develop effective, biological-based solutions to address the ecosystem disruption caused by invasion of the chestnut blight fungus (Cryphonectria parasitica) and death of billions of American chestnut trees (Castanea dentata).
In the shorter term, we continue to build on our understanding of two biological approaches to the control of chestnut blight disease: use of biocontrol agents to control the fungus populations, and tree breeding to improve the fitness of the trees. Efforts to understand the molecular, organismal, and ecological basis of C. parasitica infection, and the critical role of hypoviruses and defective mitochondria, are essential for successful development and deployment of IPM programs. NE-140 is further developing hypovirulent strains of C. parasitica for use in IPM programs to protect timber and orchard chestnut trees, and seeking environmentally safe techniques to control pathogens such as Phytophthora root rot and pests such as the oriental chestnut gall wasp.
Currently NE-140 is: 1. incorporating, by standard breeding and transgenic approaches, chestnut blight resistance and gall wasp tolerance into chestnut for planting in forests and orchards 2. working to conserve native Castanea population diversity for future use in forests and orchards, including working with the National Plant Germplasm System of the USDA to establish germplasm repositories in California and Missouri 3. studying the ecology and cultural requirements of chestnut in nurseries and natural settings, leading to the reintroduction of chestnut into the forest 4. testing new chestnut cultivars, systematically documenting information on existing cultivars, establishing germplasm repositories for cultivar material, and testing new orchard management methods to enable U.S. growers to compete against U.S. fresh market imports
NE 140 has broadened its membership over the past five years to reflect the increasing breadth of chestnut research activity and bring additional research expertise to bear. The members of this group are strongly dependent on each other for the success of their individual projects. Working collaboratively, they seek to reverse the effects of devastating pathogens, find ways of dealing with new imported pests, and conduct research in support of the nascent horticultural chestnut industry.
The technical feasibility of the research. NE-140 is one of the nation's most productive research groups. Over the last five years (1998-2002), publications included 86 research papers, book chapters, reviews, and popular articles (plus an additional 169 publications from 1993-1997). Due to the growing interest in chestnut throughout the U.S., NE-140 also has expanded the breadth of institutions, researchers, and research sites beyond the Northeast region to more effectively address the multiple issues and foci related to chestnut.
The advantages for doing the work as a multistate effort. The scope, breadth, and magnitude of the effort required to combat chestnut blight disease, address other critical pathogen and pest problems, understand ecosystem processes involved in reestablishment of chestnut in the natural forest, and develop chestnut as a new orchard crop alternative for growers are far beyond the capability of any individual researcher or institution. Combined talents in molecular biology, genomics, virology, entomology, pathology, ecology, and horticulture, available across many U.S. institutions and use of numerous test sites throughout the Northeast and into the Midwest (plus a site in California) are required to effectively address the issues facing NE-140.
What the likely impacts will be from successfully completing the work. Successful completion of NE 140 goals will lead to a number of positive impacts including: reestablishment of timber chestnut trees in eastern N. America; improved stand diversity of eastern hardwood forests; successful reintroduction of chestnuts into xeric sites formerly populated by oak species decimated by gypsy moth infestations, in order to provide mast and wood products; establishment of productive nut-bearing chestnut trees in both eastern and western N. A. where a new industry is rapidly developing; and, improved economic opportunities for chestnut growers in local and international markets.
Related, Current, and Previous Work:
The total decimation of the American chestnut by chestnut blight disease had significant biological and economic impacts. During the first two decades of the 20th Century, exhaustive attempts were made to control the disease. When it was apparent that all control attempts were unsuccessful, several breeding programs were initiated. The goal of these breeding programs was to develop blight resistant American chestnut. The USDA initiated one of the first resistance breeding programs in the 1920's. Unfortunately, this program was discontinued in 1960, as their efforts in breeding a blight-resistant American chestnut were unsuccessful. Hope was renewed when C. parasitica isolates with reduced virulence (hypovirulent) were discovered in Europe.The phenomenon of hypovirulence results in a lowered pathogenicity of the fungal pathogen, allowing the tree to utilize its natural defense mechanisms and maintain vigor. Cankers are not eliminated, but the tree can produce adequate callus tissue and wall off the infection. This discovery reawakened the scientific community to the long-standing issue of chestnut blight disease. Modern scientific investigative techniques could be applied to both the fungal pathogen and the tree host. Two additional breeding institutions began breeding chestnut trees for blight resistance using modern genetic principals: The University of Tennessee-Knoxville and The American Chestnut Foundation in Meadowview, VA. These two programs complimented the one at The Connecticut Agricultural Experiment Station (CAES), the only program that has continued uninterrupted since its initiation by Arthur Graves in 1930. The large collection of species and chestnut hybrids, maintained over the years by CAES, are available to all who need breeding material. When it became clear that the oriental chestnut gall wasp (Dryocosmus kuriphilus), introduced into GA in 1974, was spreading northward, the breeding programs discussed ways to incorporate screening for gall wasp resistance into their plans.
NE-140 had it beginnings in 1978, when approximately 200 people attended a symposium held in Morgantown, WV at West Virginia University. There were 34 papers given, and the ensuing discussion convinced the group that a Regional Research Committee was needed. In 1982, five experiment station scientists agreed to participate. Within a few short years, the committee grew to include 13 experiment stations and other participating academic institutions and governmental units. As a result of this groups activities, the following events have occurred:
- CT has improved the records of holdings and continues to maintain the finest collection of species and hybrids of chestnut in the world (6). CT imported hypovirulent strains from Dr. J. Grente in France (who first described the cytoplasmically-transmissible disease of the fungus) and demonstrated that these hypovirus-infected strains could control chestnut blight cankers on American chestnut trees (8). CT described a genetic system of vegetative compatibility in the blight fungus that prevented strain fusion and the transmission of hypovirulence viruses (1, 2, 3).
- NY (Cornell) confirmed, by making genetic tests of Asian and American populations, that the blight fungus entered the US from Japan (57).
- MI described the spread of American hypovirulence viruses through the blight fungus population in a stand of trees planted in MI (45). MI described a type of hypovirulence determined by genes in mitochondria (56). MI presented a physical and genetic map of the mitochondrial genome of a strain of the blight fungus (13). MI, NJ, and WV described and compared three American hypovirulence viruses (29, 71).
- NJ, MD, MI, and TX spearheaded the naming of Cryphonectria hypoviruses as a new family (Hypoviridae), now recognized by the International Committee on Taxonomy of Viruses (49,50).
- MD transformed virulent strains of the blight fungus with cDNA copies of hypovirulence virus RNA genes, and produced stable, transgenic hypovirulent strains with virus genes and fungal genes together in the nucleus (21,22).
- MD, CT, and WV were granted permission from USDA/Plant Quarantine to test transgenic hypovirulent strains of the chestnut blight fungus in the forests of CT and WV. This was the first permit granted to test transgenic organisms for their ability to spread and effect biological control of a plant disease (7).
- KY, MI, TX, MD, VA, and CT have made detailed studies of enzymes produced by the blight fungus that may be related to its ability to kill chestnut trees (5, 15, 20, 23, 24, 25, 30, 31, 32, 35, 37, 38, 41, 43, 44, 52, 53, 56, 62, 70, 76).
- KY and CT have studied enzyme systems in chestnut trees that may be related to their ability to resist the blight fungus (36, 39, 40, 61, 65, 66, 67,68,69).
- VA, CT, and TN have been breeding chestnut trees and selecting progeny for resistance to chestnut blight disease and with the timber-form of American chestnut trees (4). These will be ready for release within five years.
- MS, VA, MA, and CT prepared a genetic map of chestnuts and found three molecular markers associated with resistance to chestnut blight disease (54).
- MO, TN, MI, KS, CT, and CA have initiated cultivar trials of exotic chestnuts (Chinese, Japanese, and European in origin), with the goal of developing chestnut research orchards, thereby creating a U.S. chestnut industry.
- Studies conducted by MI, MN, NY, WV, and WI on the effectiveness of two hypoviruses to affect biological control, conducted at an American chestnut stand in West Salem, WI, have shown that hypovirus spread and biological control has been most significant on trees where hypoviruses were introduced (27,51). The best surviving hypovirus is the one less debilitating to C. parasitica. Also, new vegetative compatibility groups of C. parasitica were discovered and named. A mitochondrial plasmid was found in some West Salem isolates, but its role in disease expression is as yet unknown (11).
- A MI laboratory study on the effects of specific C. parasitica vic genes on the restriction of the hypovirus transmission was confirmed by WV in a forest setting (12). Also, a study conducted by WV and NY, compared vegetative compatibility types in the US with those in Europe (26,58). NJ and WV examined and characterized a cryptic dsRNA (CHV4-SR2) found in 30% of the isolates recovered from Appalachia.
- MD reported that "transgenic hypovirulent" strains are able to transmit virus to sexual (ascospore) progeny. This novel virus transmission property is predicted to provide increased biological control potential by circumventing barriers imposed by the fungal vegetative incompatibility system (9).
- WV and VA, involved in an ongoing long-term study of hypovirus dissemination in cleared and non-cleared, cut-over areas, demonstrated that chestnut regeneration was perpetuated by continuous clearing, while hypovirus remained after repeated hypovirus introductions. MD, CT, and WV have shown that the sexual recombining ability of transgenic with virulent isolates was high when tested in a forest setting but evidence for hypovirus dissemination was lacking (9,28).
- WV and NY have collaborated on several projects related to C. parasitica population structure. C. parasitica hypoviruses that survived in WV forest settings 15 years after their release, typically were avirulent (16,17,55).
- WV conducted two experiments to examine some of the parameters associated with hypovirus transmission in the field. The influence of mycelium age on hypovirus transmission within cankers was examined using two hypoviruses, CHV1-Euro 7 and CHV3-COLI 11-1 (data not published). Canker age and time of hypovirus introduction on the development of hypoviruses in cankers was examined using the above mentioned hypoviruses (data not published).
- Work conducted in MD has resulted in the preparation of Expressed Sequence Tagged (EST) libraries with mixed mRNA populations isolated from both hypovirus-infected and uninfected cultures. cDNAs were cloned into a lZipLox system to facilitate easy recovery of insert fragments contained in the pZL1 vector background. Recovered ampicillin resistant colonies were picked into microtiter plates, catalogued, and stored at -80oC. Characterization of the libraries are in progress.
- MD has identified hypovirus encoded genes that contribute to alterations in C. parasitica colony morphology, growth characteristics, sporulation levels and the size and morphology of canker formation on chestnut tissue (18,19,59,72,73).
- Work in CA has progressed to understand the molecular interaction between C. parasitica and the hypovirus CHV-1, specifically the attenuation of virulence. This includes isolation of C. parasitica genes down-regulated by the virus at the level of transcription, and characterization of cryparin, one of the down-regulated genes shown to play a critical role in the formation of fruiting bodies on wood. The viral polymerase and its products have been characterized. Cloning, sequencing and deletion of Mf1-1 and Mf2-1, mating type specific pheromones has been accomplished. Identification of 2 extracellular and 1 intracellular laccase has occurred. Deletion of one of the extracellular laccases was shown to have no phenotypic effect. Cryparin secretion has been characterized.
- MA has found that microorganisms antagonistic to C. parasitica, such as the fungus Trichoderma atroviride and the bacterium Bacillus megaterium, show biocontrol potential (47,48,74). MA research has demonstrated that both B. megaterium and T. atroviride contain genes for antagonism against C. parasitica . Similar genes for antagonism to C. parastica, however, may also exist in other fungi and bacteria on the bark of the American chestnut, which may play a key role in the microbial ecology of the chestnut blight disease.
- A symposium on hypovirus deployment was presented at the annual American Phytopathological Society meeting in Milwaukee, WI (July, 2002). MI, NY (Cornell), TN, and WV were involved in the symposium.
Objectives
- To improve chestnut trees for reestablishment in forest ecosystems, and chestnut cultivars for nut production by selection, breeding, and marketing, and determine the cultural criteria of all chestnuts for successful production in nurseries, orchards, and/or natural settings.
- To evaluate and integrate multiple approaches for the biological control of the chestnut blight fungus and other pathogens and pests that threaten chestnut, by investigating host/pathogen/parasite relationships from the molecular to the ecological level.
