Journal of Forestry ¥ D ecember 20014

G enetic engineering, commonlycalled genetic modification(GM ) in much of the world, is

the use of recombinant D N A and asex-ual gene transfer methods to breedmore productive or pest-resistantcrops. I t has been the subject of con-siderable controversy, with concernsraised from biological, socioeconomic,political, and ethical perspectives.Some of the issues are similar to thoseraised by the use of molecular biologyand genetic engineering in medicine,which we see in the news headlinesdaily. H owever, genetic modificationin agriculture and forestry raises envi-ronmental issues as well.

G M crops, mainly herbicide- andpest-resistant varieties of soybeans,maize, or cotton, have been vigorouslyadopted by farmers in North Americabecause they are easy to manage andthey improve yields, reduce costs, or re-duce pesticide ecotoxicity (C arpenter

and Gianessi 2001). H owever, the con-troversy, primarily embodied in regula-tory barriers to trade of GM crops withEurope and Japan, has slowed theiradoption considerably in recent years.

I f G M trees are used in forestry inthe near future, they are likely to occurprimarily in intensively managed envi-ronments, such as urban forests orplantations. In urban forestry, geneticmodification is expected to help treesadapt to the stresses and special de-mands of human-dominated systems.Examples would be trees that are moretolerant of heavy metals or other pollu-tants, resist urban pests or diseases,grow slower, or do not produce fruitswhen these create hazards in street en-vironments (Brunner et al. 1998).

Plantations, although very differentfrom natural forests in structure andfunction, are considered part of thespectrum of methods in sustainableforest management (R omm 1994).

Plantations can relieve pressure on nat-ural forests for exploitation and can beof great social value by supplying com-munity and industrial wood needs andfueling economic development. T heenvironmental role of plantations isrecognized by the Forest StewardshipC ouncil (FSC ), an international bodyfor certification of sustainably man-aged forests. FSC Principle 10 statesthat plantations should Òcomplementthe management of, reduce pressureson, and promote the restoration andconservation of natural forestsÓ (FSC2001).

FSC has certiÞed some of the mostintensively managed plantations in theworld, including poplar plantationsand the intensive pine and eucalyptplantations of the Southern H emi-sphere. Although many environmentalmitigations are built into these certiÞedplantation systems, within the areasdedicated to wood production theyfunction as tree farms. Such intensiveplantation systems often use highlybred genotypes, possibly including ex-otic species, hybrids, and clones, aswell as many other forms of intensivesilvicultural management. I t is in thecontext of these biointensive systemsthat the additional expense of G Mtrees is likely to be worthwhile.

H owever, FSC currently prohibitsall uses of GM trees, and is the only cer-tification system to have done so

S teven H. S trauss , Malcolm M. C ampbell, S imon N. Pryor,Peter C oventry, and J eff B urley

Genetic engineering, also called genetic modification (GM), is the isolation, recombinant mod-ification, and asexual transfer of genes. It has been banned in forest plantations certified by theForest Stewardship Council (FSC) regardless of the source of genes, traits imparted, or whetherfor research or commercial use. We review the methods and goals of tree genetic engineeringresearch and argue that FSCÕs ban on research is counterproductive because it makes it diffi-cult for certified companies to participate in the field research needed to assess the value andbiosafety of GM trees. Genetic modification could be important for translating new discover-ies about tree genomes into improved growth, quality, sustainability, and pest resistance.

Keywords: biotechnology; entomology and pathology; ethics; genetics; silviculture

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Plantation C ertiÞcation & Genetic Engineering

FSC Õs Ban on Research Is C ounterproductive

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(Coventry 2001; Strauss et al. 2001a).FSC Principle 6.8 simply states the “useof genetically modified organisms shallbe prohibited” (FSC 2001). This policystands in stark contrast to their dis-criminating, but not exclusionary, al-lowance of the use of chemicals, includ-ing pesticides, exotic biological controlagents, and exotic tree species or popu-lations. Like genetic modification, all ofthese practices can have undesirable en-vironmental consequences, includingirreversibility, that are accepted or miti-gated to the extent feasible. These prac-tices are allowed because it is felt thatthey have undergone sufficient researchand their environmental or economicbenefits outweigh their risks.

In 1999 FSC briefly outlined its rea-sons for concern over genetic modifica-tion (Strauss et al. 2001a). However,none of the problems raised appear in-soluble given adequate research, selec-tive deployment, and mitigation whenneeded. With the rapid growth of FSCcertification—now at about 22 millionhectares worldwide—this prohibitionon research will make it increasinglydifficult for industries to participate inresearch with GM trees and thus posesa significant obstacle to answering thevery questions about GM trees thatFSC has raised. There are many waysto conduct field research on GM treeswith a high degree of environmentalsafety, including harvest of trees priorto flowering (Strauss et al. 2001a). Thisconcept was supported by a unani-mous resolution of the IUFRO Sectionon Molecular Biology of Forest Trees(2001) at its international meeting inJuly 2001; the section requested thatFSC reconsider its blanket exclusion ofGM trees in research.

GM versus Conventional BreedingGenetic modification is similar to

conventional breeding in that it seekschanges in the genetic constitution oftrees to make them more productiveunder the environmental conditions ofplantation management or urban for-estry. However, in contrast to tradi-tional breeding where natural variationis selected based on plant phenotype(appearance), in genetic modification

the genes that control desired traits arephysically isolated, their DNA se-quences determined, and the genesthen modified and reintroduced via anasexual process, usually in a petri dish.Genetic modification therefore reliesheavily on the nascent but rapidlygrowing knowledge of genes andgenomes. It is also concerned withtraits that can be usefully modifiedwith one or a few genes. Because of theconservation of the genetic code, andbecause at this fundamental geneticlevel genes from different organismsshow striking similarities in their struc-ture and function, genes that were iso-lated from other organisms can beused. Transfer of one or a few genesfrom the tens of thousands that makeup an organism do not effectively makethem hybrids or chimeras—althoughFrankensteinian metaphors do appearto help promote anti-GM campaignsand sell newspapers.

Traditional breeding, in contrast,tends to focus on complex traits, suchas adaptation to environment, that de-pend on the interactions of large num-bers of genes. Genetic modification istherefore not a replacement for tradi-tional breeding but a way to solve spe-cific problems or to add value to theproducts of an advanced conventionalbreeding program. It is most easily em-ployed when breeding has proceededto the stage of clonal propagation, suchas in poplars, eucalypts, and someconifers, where it can be applied to al-ready commercially valuable clones.

Genetic modification seeks both toadd new traits not available in the na-tive gene pool, such as the new forms ofherbicide or pest resistance seen in agri-cultural crops, and to modify nativegenes in specific ways to increase pro-ductivity or improve management effi-ciency. The goals of adding new traitsare to reduce pest control costs, reduceor avoid pesticide use, improve envi-ronmental profiles of herbicides used,or facilitate low tillage systems to re-duce erosion and soil damage. At leastone of these benefits has been realized,sometimes to a striking degree, with thefirst generation of GM agriculturalcrops (Carpenter and Gianessi 2001).

As for modifications to native genes,one application is to alter genes thatcontrol xylem development so that thewood produced is better suited to spe-cialized end uses, such as pulping orbioenergy production, allowing higheryields or more economical or environ-mentally benign processing. Verypromising results have already beendemonstrated from studies of GMpoplars (Dinus et al. 2001). In anotherexample, fertility can be reduced bymodifying native genes critical to re-production. This application is ex-pected to increase wood productivity,aid production of hybrids in bisexualspecies (male sterility), and reduce thespread of introduced genes to wildpopulations (Strauss et al. 1995). Inaddition, when trees are threatened bypests for which native resistance is rareor lacking (e.g., Dutch elm disease andchestnut blight), genetic modificationshould help to mobilize genes from re-lated species, including other speciesand genera, that have developed resis-tance due to long-standing associationwith the exotic pests. This capabilitywill grow as genomes are bettermapped and understood (Adams et al.,in press).

In addition to knowledge of genesand genomes, the use of genetic modi-fication depends on asexual transfer ofgenes into cells and recovery of healthytrees. These methods are well advancedfor poplar (aspens and cottonwoods),sweetgum, and a few other species. Inpoplars, the majority of field trials con-ducted around the world have shownthat GM trees grow well and expresstheir new traits with stability (Pilate etal. 1997; Strauss et al. 2001b). Formost tree species, however, substantialadditional research is needed.

Like the products of conventionalbreeding, newly produced GM treesmust be vigorously field-tested to iden-tify the most desirable genotypes and toensure that they provide the requiredvalue, adaptation, and environmentalsafety over a range of conditions. Thecurrent regulatory system in the UnitedStates, where GM trees are under thecontrol of the US Department of Agri-culture and the Environmental Protec-

5December 2001 • Journal of Forestry

tion Agency, give legal strength to thisbasic business practice and principle ofgood forest stewardship.

Politics versus ScienceOne of the unfortunate conse-

quences of this controversy is the ten-dency to treat all of genetic modifica-tion as a unitary group that is eithergood or bad. In debate, the argumentsoften shade from biological to ideolog-ical, depending on the worldview ofthe participant. Those against intensivemanagement for wood production,who feel genetic modification is unac-ceptably unnatural or who object tothe highly patent-intensive and thuscorporate role in genetic modification,tend to dislike it. Those who believethat growing more wood on less land isan important environmental as well aseconomic goal, and who accept a con-tinuing major role for technology andlarge corporations in forestry and agri-culture, tend to favor it. Professionaland public opinions can thus becomehighly polarized (Priest 2000).

However, the biological reality ofgenetic modification does not supportsuch black-and-white judgments(Strauss et al. 1999). Genes differ dra-matically depending on what they doand how they are modified. The healthof GM trees varies widely dependingon the species, genes, and gene transfer

method used. The consequences andsafety of the new traits depend on thesilvicultural systems in which the treesare managed. Even at the social level,ethical judgments of desirability willdepend on assessments of benefit ver-sus risk, which vary with socioeco-nomic context.

Perhaps the most important conse-quence of FSC’s ban on GM research isthat it tends to foster the view that allgenetic modification is dangerous tothe environment. In contrast, leadingscientists, including the National Acad-emy of Sciences and the Ecological So-ciety of America, long ago stated, andrecently reconfirmed, that the newtraits imparted by genetic modification,not the method, should be the focus ofbiosafety analysis (Tiedje et al. 1989;National Research Council 2000).

The polarization of views about ge-netic modification can have seriousconsequences. It gives license to strin-gent regulatory regimes based onmethod and may encourage broadscaleattacks against all GM research and de-velopment, even when research is ori-ented toward biosafety methods (Kaiser2001). The May 2001 arson attackagainst the University of WashingtonProgram in Urban Horticulture, whichdestroyed much of an entire buildingand the work of diverse researchers (andwhich easily could have taken lives) is a

particularly tragic example of wherethis polarization can lead.

Moving ForwardIt is not surprising that the qualita-

tive genetic changes allowed by geneticmodification can have substantialvalue. The biological demands on plan-tation trees, particularly those managedfor specific fiber, energy, chemical, orsolid wood products, are very differentfrom the demands on wild trees. Forexample, changes in wood chemistrythat might have deleterious effects onlong-lived, wild trees are likely to betolerable in highly managed, short ro-tation systems. Even minor changes infeedstock quality can yield tremendouseconomic and environmental benefits(Dinus et al. 2001). Such changes arealso likely to domesticate rather thaninvigorate trees, making them less ableto compete in wild environments.

Precisely what kinds and degrees ofalterations to woody quality, as well asother traits, are biologically acceptableand deliver significant value can onlybe determined by research, particularlyfield trials. However, because of thecosts, regulatory demands, proprietaryissues, and long-term nature of fieldtrials, industry participation is usuallyrequired. Therefore, as more compa-nies become certified by FSC, con-ducting the needed research to evaluateGM trees will become increasinglyproblematic.

The public, not to mention manyprofessionals, often are confused by thecomparison of GM trees to infamousinvasive exotic species such as kudzuvines and gypsy moths. GM plants areindeed “exotic” in the sense that theydeliver new traits that are rare or absentin wild gene pools. However, they arenot exotic in that they are not biologicalnovelties in ecosystems. When exoticorganisms are disruptive it is generallybecause they occupy a new niche, pos-sess novel and complex adaptations,and have many new, interacting geneticnetworks not present in native species.They also generally leave behind hun-dreds of pests when they depart theirnative ecosystems. The simple, limitedgenetic alterations imparted by geneticmodification bear little resemblance tothis panoply of novelties.

6 Journal of Forestry • December 2001

An FSC-certified, intensively managed hybrid poplar plantation in eastern Oregon. Because of theiramenability to gene transfer, facile clonal propagation, and intensive silviculture and breeding,poplar fiber farms such as this one are ideal places for research and possible deployment of geneti-cally engineered trees.

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The relative precision of GM treesallows much more predictability of theirecological consequences compared toexotic species, and intensive field testingand progressive deployment allowspoorly adapted genotypes to be dis-carded. For traits whose spread outsideof plantations is undesirable, such asmight be the case for herbicide-toleranttrees, methods such as engineered infer-tility combined with vegetative propa-gation should allow the rate of spread tobe drastically reduced or avoided en-tirely. This would also reduce ecologicalimpacts from the spread of exotic andconventionally bred trees. Although thetechnology for engineered sterility sys-tems has already been demonstrated intransgenic agricultural crops, the effi-ciency and biosafety of this option re-quires field study in GM trees.

As tree genomes are better under-stood, the options for using detailed ge-netic information to more precisely tailortrees to intensive production systems aregrowing rapidly. Because of the limita-tions imposed by the long life cycle andoutbreeding system of mating for trees—which greatly slow the rate of progress orlimit breeding options—genetic modifi-cation may present an important futuremeans for making genetic improve-ments. A key avenue for progress wouldtherefore be foreclosed if biopoliticalpressures, such as are embodied in FSC’sban on genetic modification, impede sig-nificant research in this area.

Instead of slowing research, whatwould be most socially valuable wouldbe for FSC and other certification sys-tems to help ensure that GM trees arestudied and deployed responsibly. Thiswould be most important in develop-ing countries, which often do not havesubstantial regulation and associatedinfrastructure. The stewardship obliga-tions that should accompany geneticmodification are very similar to thoseinvolved with use of other technologiessuch as chemicals and exotic or bio-control organisms and could be readilyadopted as parts of certification assess-ments (Strauss et al. 2001a).

Ethical ChallengesBiotechnology in medicine and agri-

culture presents a variety of biologicaland social complexities. The potential

benefits from applying detailed geneticknowledge are obviously great, butthere is much social debate about whatconstitutes sustainable and ethical ap-plication in various sectors. In forestry,the obvious near-term applications arein highly intensive systems, such asplantation and urban forestry. How-ever, even these applications requireconsiderable research to define safe andproductive uses. Because of the longtime period required for tree research, itseems wise to conduct research in par-allel with the social debate about safeand appropriate applications of geneticmodification in forestry. Demonstra-tions of benefit and safety ultimatelywill be required for ethical judgmentsof desirability in specific circumstances(Thompson and Strauss 2000).

We suggest that certification sys-tems, including FSC, promote (or atthe very least not prohibit) research toidentify productive and safe uses of ge-netic modification in intensive forestmanagement. With the wealth of sci-entific possibility provided by the ge-nomics revolution, the great uncer-tainty about stresses on future forestsand societies, and the diversity of needsand management problems, prohibit-ing all applications of genetic modifi-cation, and particularly field research,is neither wise nor precautionary.

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Steven H. Strauss (steve.strauss@orst.edu)is professor, Department of Forest Science,Oregon State University, RichardsonHall, Corvallis, OR 97331-5752; PeterCoventry is associate professional officer,UK Department for International Devel-opment, Pará, Brazil; Malcolm M.Campbell and Simon N. Pryor are lec-turers and Jeff Burley is director, OxfordForestry Institute, Department of PlantSciences, Oxford, UK. This article waspartially supported by a grant from theScottish Forestry Trust.