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Technological Advances in Temperate Hardwood Tree Improvement Including Breeding and Molecular Marker Applications

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Technological Advances in Temperate Hardwood Tree Improvement Including Breeding and Molecular Marker Applications In Vitro Cell.Dev.Biol.—Plant (2007) 43:283–303 DOI 10.1007/s11627-007-9026-9 INVITED REVIEW Technological advances in temperate hardwood tree improvement including breeding and molecular marker applications Paula M. Pijut & Keith E. Woeste & G. Vengadesan & Charles H. Michler Received: 20 September 2006 /Accepted: 8 January 2007 / Published online: 6 July 2007 / Editor: P. Lakshmanan # The Society for In Vitro Biology 2007 Abstract Hardwood forests and plantations are an impor- Genetic modification of hardwood tree species could tant economic resource for the forest products industry potentially produce trees with herbicide tolerance, disease worldwide and to the international trade of lumber and logs. and pest resistance, improved wood quality, and reproduc- Hardwood trees are also planted for ecological reasons, for tive manipulations for commercial plantations. This review example, wildlife habitat, native woodland restoration, and concentrates on recent advances in conventional breeding riparian buffers. The demand for quality hardwood from and selection, molecular marker application, in vitro tree plantations will continue to rise as the worldwide culture, and genetic transformation, and discusses the consumption of forest products increases. Tree improve- future challenges and opportunities for valuable temperate ment of temperate hardwoods has lagged behind that of (or “fine”) hardwood tree improvement. coniferous species and hardwoods of the genera Populus and Eucalyptus. The development of marker systems has Keywords Clonal propagation . Cryopreservation . become an almost necessary complement to the classical Forest genetics . Genetic transformation . Organogenesis . breeding and improvement of hardwood tree populations Plantation forestry. Regeneration . Somatic embryogenesis for superior growth, form, and timber characteristics. Molecular markers are especially valuable for determining the reproductive biology and population structure of natural Introduction forests and plantations, and the identity of genes affecting quantitative traits. Clonal reproduction of commercially Hardwood forests and plantations in North America, important hardwood tree species provides improved plant- Europe, and other parts of the world contain a wide range ing stock for use in progeny testing and production forestry. of temperate tree species that are an important resource for Development of in vitro and conventional vegetative the forest products industry and to the foreign trade of propagation methods allows mass production of clones of lumber and logs. In addition to timber, sawlog, and veneer mature, elite genotypes or genetically improved genotypes. log production, hardwood trees are also planted for wildlife habitat, native woodland restoration, riparian buffers, erosion control, windbreaks, conservation, and P. M. Pijut (*) : K. E. Woeste : C. H. Michler Hardwood Tree Improvement and Regeneration Center (HTIRC), watershed protection. Some of the more valuable hard- Northern Research Station, USDA Forest Service, woods include alder (Alnus spp.), ash (Fraxinus spp.), 715 West State Street, West Lafayette, basswood (Tilia spp.), beech (Fagus spp.), birch (Betula Indiana 47907, USA spp.), black locust (Robinia pseudoacacia), black cherry e-mail: [email protected] (Prunus serotina), chestnut (Castanea spp.), elm (Ulmus spp.), gum (Liquidambar styraciflua), hackberry (Celtis G. Vengadesan occidentalis), hard (and soft) maples (Acer spp.), hickory Deparment of Forestry and Natural Resources, HTIRC, and pecan (Carya spp.), oak (Quercus spp.), sassafras Purdue University, 715 West State Street, West Lafayette, (Sassafras albidum), sycamore (Platanus spp.), walnut Indiana 47907, USA (Juglans spp.), black willow (Salix nigra), and yellow 284 PIJUT ET AL. poplar (Liriodendron tulipifera). The market for these for commercial plantations is also a major aspect of a tree species can be very high because of the special appearance improvement program. Development of an effective gene (e.g., grain, figure, texture, and color) or technical proper- transfer and efficient in vitro regeneration system for each ties (e.g., strength, durability, and good machining proper- hardwood species, that can be easily adapted for many ties) compared to lesser quality hardwoods used for fuel or genotypes, will facilitate the production of genetically pulp. These species are utilized in the manufacture of improved temperate hardwood trees. residential and commercial structures and furnishings Plantation forests and the role biotechnology can play (architectural millwork, cabinets, doors, flooring, furniture, has been reviewed (Fenning and Gershenzon 2002). Other moldings, paneling, shutters, siding, and windows) and recent reviews have focused on innovative technologies other specialty products (barrel staves, baseball bats, that provide the basis for acceleration in forest tree billiard cues, boat interiors, butcher blocks, carvings, improvement (Nehra et al. 2005) and in vitro propagation, caskets, crates, gun stocks, hockey sticks, kitchen utensils, gene transfer, and genomics for a sample of hardwood ladders, musical instruments, oars, skis, tool handles, toys, timber and pulp species (Merkle and Nairn 2005). Because Venetian blinds, and woodenware). Populus and Eucalyptus biotechnology has been recently Tree improvement of temperate hardwoods has been reviewed, we will limit our review to the hardwoods more limited compared to that of coniferous species and previously mentioned. This review concentrates on con- hardwoods of the genera Populus and Eucalyptus (Merkle ventional breeding and selection, molecular marker appli- and Nairn 2005). The demand for hardwood from tree cation, in vitro culture, genetic transformation, and future plantations will continue to rise as the worldwide consump- challenges and opportunities in valuable temperate hard- tion of forest products increases, and the environmental, wood tree improvement. A comprehensive review of the commercial, and political pressures of restricting logging of literature is impossible, but hopefully we have captured or high-quality trees from natural forests also increases (for highlighted many important species and research. example, see references and working papers on forest plantations cited in FAO 2001). Considerable effort has been exerted over the last 40 yr in conventional tree Conventional Tree Improvement improvement programs through breeding and selection, and strategies for breeding and tree improvement of temperate The conventional improvement of hardwoods has always hardwoods have been developed (Burley and Kanowski lagged behind that of conifers. Deciduous hardwood 2005; Michler et al. 2005). The long generation and species that are used for fiber or horticulture, and those reproductive cycle, difficulty in conducting controlled polli- with congeners used by those industries, have benefited nations, intermittent or scarce seed crops, and seed recalci- from a research crossover effect that mostly informs trance of hardwood trees are some of the limitations imposed biotechnological methods. Conventional breeding still on conventional tree breeding programs (Lantz 2007). Forest relies on a mainstay of provenance trials to evaluate local geneticists are interested in developing populations with adaptation, phenotypic selection to identify potentially disease resistance, superior growth, form, and timber superior parents, progeny trials to evaluate those parents, characteristics, including straighter boles and reduced and seed orchards for the production of adapted, improved branching. Molecular marker development would be seed. A web resource that links to many of the tree useful in determining the genetic quality and population improvement programs (including conifers) world-wide structure of natural forests and plantations, and the (http://www.genfys.slu.se/staff/dagl/Documentations/ quantitative genes of superior trees. Clonal reproduction OrganisationLinks.htm) is maintained by Dag Lindgren at of commercially important hardwood tree species is also SLU, Umeå, Sweden. In Europe, recent activities of the necessary in a tree improvement program to provide British and Irish Hardwood Improvement Programme are improved planting stock for use in progeny testing and summarized in Burley (2004), including research in ash, for production forestry. In vitro and conventional vegeta- silver birch, wild cherry, two species of oak, sweet chestnut, tive propagation methods will be required to produce sycamore (Acer), and walnut (two species). Recent research clones of mature, elite genotypes or genetically improved in Europe and elsewhere has focused at least as much on genotypes. Many economically important hardwood tree conservation of genetic resources as on the development species have a low genetic or physiological capacity for (breeding) of resources (Eriksson 2001; Xie et al. 2002; adventitious root formation, and are considered recalci- Karagöz 2003; Hosius et al. 2006), but these two activities trant to routine, commercial-scale vegetative propagation. are closely related and interdependent. Reviews focused Genetic modification of hardwood tree species to produce specifically on tree improvement in particular regions or in trees with herbicide tolerance, disease and pest resistance, particular species are also available (Koski and Rousi improved wood quality, and reproductive manipulations 2005). In the US, summaries of hardwood breeding TEMPERATE HARDWOOD TREE IMPROVEMENT 285 research
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