6. REFERENCES

Adam, D. 2003. Transgenic crop trial’s gene flow turns weeds into wimps. Nature 421:462.

Anonymous. 2003. Missing the big picture. Nature 421:675.

Arntzen, C.J., Coghlan, A., Johnson, B., Peacock, J., Rodemeyer, M. 2003. GM crops: science, politics and communication. Nature Reviews Genetics 4:839-843.

Campbell, M.M., Brunner, A.M., Jones, H.M., Strauss, S.H. 2003. Forestry’s fertile crescent: the application of biotechnology to forest trees. Forest Biotechnology 1:141-154.

Crawley, M.J., Brown, S.L., Hails, R.S., Kohn, D.D., Rees, M. 2001. Transgenic crops in natural habitats. Nature 409:682-683.

Dalton, R. 2003. Superweed study falters as seed firms deny access to transgene. Nature 419:655.

DiFazio, S.P., Leonardi, S., Cheng, S., Strauss, S.H. 1999. Assessing potential risks of transgene escape from fiber plantations. Lutman, P., ed. Proc. Symp. Gene flow and agriculture: relevance for transgenic crops. No. 72. British Crop Protection Council, Farnham, UK.

Ellis, D., Mellan, R., Pilate, G., Skinner, J.S. 2001. Transgenic trees: where are we now? pp. 113-123, Strauss, S.H., Bradshaw, H.D., eds. Proc. first international symp. on ecological and societal aspects of transgenic plantations. College of Forestry, Oregon State Univ., Corvallis, OR.

FAO. 2001. Biotechnology in the forest sector. http://www.fao.org/biotech/sector5.asp

FAO. 2002. Conference 7. Summary Document, FAO electronic conference on Gene flow from GM to non-GM populations in the crop, forestry, animal and fishery sector. http://www.fao.org/biotech/logs/C7/summary.htm

Forest Stewardship Council (FSC). 1996, revised 2000. FSC Principles and Criteria. Document 1.2, Principle 6.

Gartland, K.M.A., Kellison, R.C., Fenning, T.M. 2002. Forest biotechnology and Europe’s forests of the future. Proc. forest biotechnology in Europe: impending barriers, policy and implications. Edinburgh.

Government of Canada. 1985 (revised 2003). Seeds Act. c. S-8. http://laws.justice.gc.ca/en/S-8/98571.html

Haines, R.J. 1994. Biotechnology in forest tree improvement: research directions and priorities. Forestry Paper 118, UN FAO, Rome.

Hancock, J.F., Hokanson, K.E. 2001. Invasiveness of transgenic vs. exotic plant species: how useful is the analogy? pp. 187-192, Strauss, S.H., Bradshaw, H.D., eds. Proc. first international symp. on ecological and societal aspects of transgenic plantations. College of Forestry, Oregon State Univ., Corvallis, OR.

Heron, D.S., Kough, J.L. 2001. Regulation of transgenic plants in the United States. Pp. 101-104, , Strauss, S.H., Bradshaw, H.D., eds. Proc. first international symp. on ecological and societal aspects of transgenic plantations. College of Forestry, Oregon State Univ., Corvallis, OR.

Industry Canada - Life Sciences Branch. 2001. Biotechnology in the forestry sector. Natural Resources Canada. http://strategis.ic.gc.ca/bo01532e.html

International Board for Plant Genetic Resources (IBPGR, comp.). 1991. Elsevier's Dictionary of Plant Genetic Resources. IBPGR Secretariat, Rome, Italy.

Iowa State University. 1994. Principles of Biotechnology. (Bio-1). North Central Regional Extension Publication, Cooperative Extension Service, Iowa State University, Ames, IA http://www.nal.usda.gov/bic/Education_res/iastate.info/bio1.html

Johnson, B., Kirby, K. 2001. Potential impacts of genetically modified trees on biodiversity of forestry plantations: a global perspective. pp. 176-186, Strauss, S.H., Bradshaw, H.D., eds. Proc. first international symp. on ecological and societal aspects of transgenic plantations. College of Forestry, Oregon State Univ., Corvallis, OR.

Krutovskii, K.V., Neale, D.B. 2001. Forest genomics for conserving adaptive genetic diversity. Forest Genetics Working Paper FGR/3. Forest Resources Development Service, Forest Resources Division, FAO, Rome.

Libby, W.J. 1973. Domestication strategies for forest trees. Canadian Journal of Forest Research 3:265-276.

Owusu, R.A. 1999. GM Technology in the forest sector: a scoping study for WWF. http://www.panda.org/resources/publications/forest/gm/gm-risk.html

Pew Initiative on Food and Biotechnology. 2001. Biotech branches out: a look at the opportunities and impacts of forest biotechnology. Proc. workshop, Pew Initiative on Food and Biotechnology, Society of American Foresters, Ecological Society of America. Atlanta, GA. 28 pp.

Pilate, G., Guiney, E., Holt, K., Petit-Conil, M., Lapierre, C., Leplé, J.-C., Pollet, B., Mila, I., Webster, E.A., Marstorp, H.G., Hopkins, D.W., Jouanin, L., Boerjan, W., Schuch, W., Cornu, D., Halpin, C. 2002. Field and pulping performances of transgenic trees with altered lignification. Nature Biotechnology 20:607-612.

Stone, G.D. 2002. “Both Sides Now: Fallacies in the Genetic Modification Wars, Implications for Developing Countries, and Anthropological Perspectives,” Current Anthropology 43(4), Aug.-Oct. 2002.

Ritland, C., Ritland, K. 2000. DNA-fragment markers in plants. pp. 208-234, Baker, A.J., ed. Molecular methods in ecology. Blackwell Scientific, Oxford, UK.

Schmidt, L. 1997. Tree improvement glossary. Technical Note No. 46. Danida Forest Seed Centre, Humlebæk, Denmark.

Sedjo, R.A. 1999. Biotechnology and planted forests: assessment of potential and possibilities. Discussion paper 00-06, Resources for the Future, Washington, D.C. 28 pp.

United Nations Environment Programme (UNEP). 1992. The Convention on Biological Diversity. Article 2. Environmental Law and Institutions Programme Activity Centre. United Nations, New York and Geneva.

Yanchuk, A.D. 2001. The role and implications of biotechnological tools in forestry. Unasylva 204(52):53-61.

Table 1: Anticipated contribution to and scale of impact of each broad area of biotechnology on elements of natural and artificial forest populations

Table 2: Current global applications and projected future importance and trends of forest tree biotechnology
A = developed countries; B = developing countries; on a scale of 0-3: 0 = nil, 3 = common.

Broad Technologies	Components	Current Applications		Projected Trends
		A	B
	Large databases:
Bioinformatics	Targeted DNA Sequence	3	0	The storage, retrieval, analysis, and interpretation of large amounts of biological data will cross boundaries of all broad biotechnologies. Capability and application range of this tool will continue to increase dramatically. Mining the massively increasing amounts of data at all scales, and integrated analyses and syntheses of these data will greatly increase power to detect genes and understand their functions. The databases developed for sequence and microarray data in particular must be co-ordinated regionally, nationally and internationally, and accessible to the public. Bioinformatics research requires resource-intensive, multidisciplinary teamwork, naturally leading towards more international cooperation over a range of study scales and systems. Opportunities are opening up for developing countries to become involved in these projects.
	Proteomic	2	0
	Gene Mapping & Markers	3	2
	Microarray	2	0
	Phenotypic	3	3
	Integrated Applications	1	0
Diversity Measurement	mtDNA	2	0	The use of molecular markers for studying natural and artificial forest tree populations has undergone unprecedented expansion due to the vast array of population genetics applications they have enabled. These topics include measurement of genetic diversity within and among populations, comparisons among taxa, historical reconstruction and prediction of species’ range shifts, gene flow, assessment of natural and artificial (e.g., seed orchard) population mating system parameters, introgression and hybridization. Markers have also been used to evaluate the impacts of domestication and silviculture. Genomic and QTL mapping have recently expanded due to the development of unlimited numbers of markers. Anticipated development cost reductions for SNPs will trigger an increase in their use in all forest genetics resources applications.
	cpDNA	2	0
	RAPD, AFLP, RFLP	3	2
	Microsatellite (SSR)	1	0
	SNP, ESTP	3	1
Gene Discovery	Phenotypic Traits	3	3	Phenotypic and quantitative trait measurements represent the backbone of all conventional tree breeding. Their proven efficacy and ease of use has resulted in significant gains in many species worldwide.
	QTL Mapping	3	1	QTLs will likely increase in importance, particularly for important or hard to measure traits, especially those which require older material for assessments (e.g., all wood properties, disease and insect resistance). The focus is shifting from flanking region markers to markers within the actual gene or QTL of interest.
	Genome & EST Sequencing	2	0	Massive, redundant conifer genomes will restrict sequencing to a few regions of interest. EST sequencing for commercially important species will accelerate within 5 years. Whole genome sequencing has begun for Populus; the data will be freely available; other economically and ecologically important species will follow. International collaboration is important for species whose ranges cross international borders.
	Microarray Analysis	2	0	As microarrays become available for gene discovery for growth and yield, wood quality and adaptive attributes (e.g., disease and insect resistance and stress tolerance), more reliable oligo-based will likely replace cheaper clone-based arrays. The potential benefits and challenges will spur international collaboration. This new field is likely to prove most cost-effective for understanding gene function, rapid production technology development for advanced breeding programs and in high yield plantations.
	Proteomic Analysis	2	0	Notes for microarray analysis also apply here. Proteomics airms to elucidate protein variation beyond simple transcriptional regulation, including levels of expression, interactions and post-translational modification.
	Metabolomics	1	0	Still in the initial stages, metabolomics asseses the presence/absence of non-protein structural precursors of essential components in biochemical pathways.
Molecular Genetic Modification	Gene Insertion/ Sequence Modification	2	1	Inserting foreign targeted genes into tree genomes has profound potential. This method will enable cross-species gene transfer in cases where it is not possible via conventional breeding. Transformation is widely used in agriculture, but is new to forestry and has engendered major public contention and strict biosecurity protocols for testing and deployment. Some countries and organizations have restricted testing and planting genetically modified trees. Potential for gene escape into wild populations needs further study for risk assessment. Reproduction must be eliminated or postponed past rotation age. This technique could potentially improve fibre yield and quality as well as other important qualitative, quantitative and adaptive attributes.
	Gene Targeting/ Knockout	1	0	Silencing gene function is the complement of gene insertion. This technique will expand as the results of microarray and proteomic analyses accrue. Similarly, there is a broad spectrum of potential functional genomics applications and associated ethical issues.
Product Verification	Pedigree Verification	3	3	Development of larger-scale, lower cost application platforms will increase accessibility by genetic improvement programmes to this technology, expanding the prevalent uses: retroactive pedigree verification and verifying clonal identies in seed orchards. Other potential applications include determining the efficacy of seed orchard management techniques on a large scale, and consequent growth and yield determination of resulting crops.
	Quality Control/ Quality Assurance	1	0	Rapid developments in product description systems, commonly used in food manufacturing facilities, will result in increasing support for research and application of automated product verification systems. The main technology is PCR-based, e.g., DNA markers to detect mislabelled products (e.g., clones). Functional genomics is expected to yield suites of gene markers that can be used to check process efficiencies during mass production of clonal seedlings.
Cloning	Organogenesis	1	1	Micropropagation (organogenesis) and gametic or somatic embryogenesis require treating tissue explants with growth regulators to induce bud or shoot formation. Most of the shortcomings of organogenesis can be overcome using somatic embryogenesis, but methodology and success are species dependent. These methods are currently in production and cloning will be a significant element in high yield plantation forestry.
	Somatic Embryogenesis	1	0
Biosensing	Simple Biosensors	1	0	Measuring and monitoring components of clonal production will increasingly employ sensors comprising physical, chemical and molecular markers to detect specific biological processes, e.g., expression-tagged genetic markers to quantify or detect presence/absence of metabolic pathway components of interest.