2.2 A global study on the state of forest tree genetic modification

Christian Walter and Sean KillerbyNew Zealand Forest Research Institute Ltd, Rotorua, New Zealand

2.2.1 SUMMARY

The information provided in this report was resourced from practitioners in the area through a detailed questionnaire. Particular emphasis was given to the views of practitioners on how they see genetic modification contributing to forestry, and how they view future development. To complement information from the questionnaire, other publicly available information, such as that available on Internet sites, was included in the assessment.

2.2.2 INTRODUCTION

Genetic modification in forest trees is widely reported and results from field testing are beginning to appear (Pilate et al. 2002, C. Walter, unpublished). In 1999, Owusu reported that, despite efforts by national regulatory agencies to keep track of research and applications of genetic modification in forest trees, as well as attempts to set up and maintain global information systems on GMOs (genetically modified organisms) and genetically modified (GM) trees, there was no comprehensive data set recording research and applications of genetic modification in forestry (Owusu 1999). The lack of global compilation of forest genetic modification makes it difficult to gauge the current status of such a controversial issue and to predict future trends. The present report is an attempt to quantify the existing situation with regard to developments on genetic modification in forest trees, using information and data provided by technology developers. It is, however, acknowledged that this approach can provide a snapshot of the situation only, and regular updating is recommended.

2.2.3 GLOBAL STATUS OF TREE GENETIC MODIFICATION. THE QUESTIONNAIRE

A questionnaire was developed to assess the current status of tree genetic modification worldwide, and to scope the views of scientists and organizations involved in tree biotechnology on the current and future issues related to genetic modification in trees. The questionnaire was also designed to capture the views of practitioners on future developments, and on strategies required to provide scientific information to the public. A copy of the original questionnaire is available from the FAO. Responses were collected by FAO and New Zealand Forest Research Institute Ltd (NZFRI) and analysed by the authors. The following provides a detailed analysis together with an interpretation of the results. Where appropriate, tables or graphics summarize results.

Some respondents included ornamentals/fruit trees in their responses. These were included in the analysis, because it was deemed important to capture the views of individuals and organizations practising genetic modification, with tree genetic modification as understood in the widest possible sense.

2.2.3.1 Response rate

Based on the NZFRI address list, 418 questionnaires were sent out by the authors to institutions possibly involved in genetic modification of woody plants and another 105 institutions and individuals were contacted by FAO. Further, Internet lists such as ETFRN²⁸ PT and METLA²⁹ were used to distribute information, as were the IUFRO (International Union of Forestry Research Organizations) Task Force on Biotechnology, the FAO Panel of Experts on Forest Gene Resources, the EUFORGEN³⁰ national co-ordinators and colleagues at various organizations. It can be assumed that the majority of forest geneticists, breeders and molecular biologists worldwide were informed about the study and were asked to contribute. Questionnaires were available in English, French and Spanish.

A deadline of 24 May 2003 was given for receipt of questionnaires, but this was extended, mainly owing to the fact that many key organizations had not responded by that date. The latter were contacted again directly and a few more questionnaires were returned. As of
30 September 2003, a total of 49 responses was received and entered onto a spreadsheet. This represents a response rate of 11.7 percent (based on 418 questionnaires sent originally). No more responses were received as of the end of 2003.

When data from the responding individuals and organizations were compared with publicly available data, it was noticed that not all organizations conducting research or field evaluations have made their data available, although much is publicly available from websites³¹..

2.2.3.2 Respondents (Q 1.1–1.3³²: Name, address, institutional framework, Internet)

The respondents were located in 26 different countries (Table 2.2.1).

Twenty-three of the 49 respondents (47 percent) were conducting research on genetic modification in forest trees, with one other being a non-profit organization advocating appropriate use of biotechnology.

Of the 49 respondents, 24 (49 percent) were from research institutes, 14 (29 percent) from universities, six (12 percent) from private commercial companies, and two (4 percent) from non-profit organizations. Another three (6 percent) did not state what type of organization they belonged to, or else wanted to keep the nature of their organization confidential (one respondent).

Of the 23 respondents conducting research on genetic modification in forest trees, ten
(44 percent) were in research institutes, nine (39 percent) in universities, two (8.5 percent) in a private industry organization, and two (8.5 percent) in ‘other organization’ including one confidential response. The other ‘other organization’ was CHH (Carter Holt Harvey) in New Zealand, who stated that they are currently only completing work already ‘in the pipeline’.

Fifteen of the 49 respondents (31 percent) are currently publishing information relating to genetic modification on the Internet; 21 (43 percent) do not publish.

Thirteen respondents (27 percent) did not answer these questions. However, when only those organizations that work with genetic modification were analysed, the picture is different:
13 of the 23 respondents (57 percent) working with genetic modification of trees publish information about genetic modification on the Internet. Ten (43 percent) of them do not use this medium for publishing information.

Internet addresses were submitted (see Appendix 3.8.1) by all respondents answering ‘yes’, but one did not want the Internet address to be published, for reasons of confidentiality.

Table 2.2.1. Responses by country. Numbers of respondents (total) and respondents using genetic modification are listed by country

2.2.3.3 Contained research (Q 2.1: The scope of genetic modification in contained laboratory research)

Of the 23 respondents who reported work on genetic modification of forest trees,
14 (61 percent) were conducting research on Populus, nine (39 percent) on Pinus, eight
(35 percent) on Picea and four (17 percent) on Eucalyptus. Three respondents (13 percent) were working on Betula, two (9 percent) with Larix, two (9 percent) with Chamaecyparis and two (9 percent) with Ulmus. Other species being worked on, by one respondent each, were in the genera Pseudotsuga, Robinia, Paulownia, Quercus, Juglans and Tectona³³.

When asked whether the genetic modification work was related to a research or commercial project, all but two experiments were described as research. The two commercial projects were by one organization and the taxa involved were Populus nigra and a hybrid poplar. However, three respondents classified a total of four experiments as both research and commercial. The species included were in the genera Populus and Eucalyptus (twice each), perhaps indicating that there are plans to use those trees for a commercial project in the future.

Many different traits and associated genes were mentioned by respondents as targets. Evaluation of these responses is somewhat difficult since different terms were obviously used for the same trait. Also, it can be assumed that most if not all transformation events include markers, and most probably antibiotic resistance genes, but these were not always mentioned. Table 2.2.2 (Figure 2.2.1) lists the traits by the frequency with which they were cited. Names of genes (as given in the responses) were used in this evaluation to arrive at a clearer picture of what traits were actually meant, and to group some traits.

The projects associated with commercial goals were in wood properties (three), insect resistance (two) and detoxification of pollutants (one).

A total of 68 projects was referred to and four (6 percent) of them were classified as applied for (regulatory approval), 50 (74 percent) as ongoing, and four (6 percent) as both. One project each (1.5 percent) was in development and had been stopped. However, no comments were made on eight projects (12 percent).

Table 2.2.2. Traits targeted in genetic modification projects, in laboratory based experiments

Figure 2.2.1. Traits targeted in genetic modification projects, in laboratory based experiments

2.2.3.4 Field trials (Q 2.2: The scope of genetic modification in field tests)

Eleven respondents of 49 (22.5 percent) reported projects with GM trees involving field analysis and a total of 26 projects were mentioned. Of those, 20 (77 percent) were with Populus and two (8 percent) with Pinus. One project each (4 percent each) was reported for Betula, Eucalyptus, Picea and other species in other genera.

Traits/genes targeted in these projects are listed in Table 2.2.3 (Figure 2.2.2). There are significant differences in the relative ranking of traits targeted in laboratory research
(Table 2.2.2) and traits studied in field trials, in particular with regards to reproductive development.

For 22 projects, respondents provided information on the establishment of the trials. Table 2.2.4 lists the numbers of GM tree field trials established by year. Two of the 22 projects were described as ‘ongoing’, with no commencement date given. Information on size of trial and numbers of trees, where available, is given³⁴.

When asked about the issues addressed in field trials, all but five respondents who had trials in place provided answers (Table 2.2.5 and Figure 2.2.3).

Table 2.2.3. Traits/genes targeted in projects with GM trees in field trials

Figure 2.2.2. Traits/genes targeted in projects with GM trees in field trials

Table 2.2.4. Established field trials of GM trees¹

¹–: No data provided.
²1 acre ~ 0.4 ha.

Table 2.2.5. Issues addressed in field trials with GM trees (number of projects addressing a particular issue)

Figure 2.2.3. Issues addressed in field trials with GM trees (percentage of projects addressing a particular issue)

2.2.3.5 Commercial plantations (Q 2.3: Scope of genetic modification in commercial plantations)

Most respondents reported that they had no immediate plans for commercialization, one of them commenting that it would take decades of research to test whether desired traits were realized and were stable, to assess risks, and to gain public and regulatory backing for commercial release.

One respondent felt that if there were any commercial plantations of GM forest trees in the world, those responsible were very successful in keeping it secret.

However, it has been reported (Su et al. 2003) that there are such plantations and one respondent described the commercial release of two GM tree species in China:

• Populus nigra, modified with the Bt gene³⁵ for resistance to leaf-eating insects, with 80 ha of field trials on eight sites in seven provinces since 1999 and 200–300 ha of commercial planting since 2002 (over one million cuttings have been propagated).

• Hybrid poplar 741 (P. alba × [P. davidiana + P. simonii] × P. tomentosa), modified with cry1Ac³⁶ and API³⁷ for resistance to leaf-eating insects, with field trials since 2001 and commercial planting since 2003 (currently only indicative data available on area planted).

The respondent further reported that there are significant problems keeping track of the area planted, mainly because poplar is vegetatively propagated and there is confusion in nurseries as to what is genetically modified and what is not.

2.2.3.6 Traditional breeding programmes (Q 2.4: On the support for genetic modification programmes by traditional breeding programmes)

Respondents commented on the integration of their tree genetic modification projects with traditional breeding programmes. Of a total of 68 projects involving GM trees (in 23 organizations), 27 (40 percent) reported full integration with traditional breeding projects. Nine respondents (13 percent) reported that either connections or the development of an integrated approach were weak, 29 (43 percent) indicated that their tree genetic modification projects were not integrated at all with traditional breeding projects. A further respondent who represented three projects (4 percent) made the statement that there are no links seen, and no links are anticipated.

2.2.3.7 Vegetative multiplication (Q 2.5: On the support for genetic modification programmes by vegetative propagation programmes)

Comments were requested on the integration of tree genetic modification projects with vegetative propagation projects. Of a total of 68 projects involving GM trees
(in 23 organizations), 23 (34 percent) reported full integration with clonal multiplication projects. Seventeen (25 percent) respondents reported weak connections or connections in development, while 21 (31 percent) had no integration with such programmes. Seven
(10 percent) of the respondents did not offer any information under this heading.

2.2.3.8 Other research (Q 2.6: The scope of genetic modification projects underpinning other research goals)

Under this question, respondents were asked about research projects that use genetic modification technology but do not anticipate deploying GM trees in commercial forestry. Of the 23 organizations using tree genetic modification, 15 (65 percent) reported underpinning research, seven (30 percent) answered ‘no’ and one (5 percent) did not respond. Of those organizations that did not use genetic modification for trees (27 out of 49 respondents, or
55 percent), 17 (63 percent) did not respond to this question. Eight of them (30 percent) reported no non-tree research in genetic modification, while two (7 percent) reported such research.

A total of 17 out of 49 respondents (35 percent) reported underpinning research and 12 gave more detailed descriptions of such research. The key areas were:

• testing gene function

• environmental impact

• MAS, QTL detection, genetic maps

• microarray and EST mining for genes

• flowering-related genes/cone production

• evaluation of safety, risk assessment, consumer protection

• fibre-development genes/wood formation

• biotic and abiotic stresses

• genomics and proteomics

• physiological and hormonal studies

2.2.3.9 The regulatory framework (Q 3.1: What is the regulatory framework covering genetic modification work in the respective country?)

Respondents provided information on which agency in their country was responsible for regulating genetic modification work in contained laboratories, field trials and commercial release. This information is summarized in Appendix 3.8.1. Respondents were from 15 countries and indicated whether the respective authority was involved in pre-risk assessment (13 countries), surveillance, monitoring and quarantine (12 countries), and management of failure, redress and control (12 countries). For our analysis (Table 2.2.6), only respondents involved in some type of research in genetic modification were included. Note that there are some discrepancies in responses from the same countries, probably due to misunderstandings with the questions, or different viewpoints. The data provided in Table 2.2.6 should therefore be taken with caution.

2.2.3.10 Pending applications (Q 3.2)

The aim of this question was to understand the immediate future of GM tree testing. Respondents listed a total of 12 projects pending. This includes development in containment, field testing of GM trees and release of GM trees. Note that some countries distinguish between field tests and release, whereas others do not and consider both as a release. Table 2.2.7 provides details of 12 projects.

Table 2.2.6. Involvement of regulatory agencies in pre-risk assessments of genetic modification projects, surveillance, monitoring and quarantine (S/M/Q) and management of failure, redress, control (M/R/C), by country¹.

¹–: indicates no answer.

Table 2.2.7. GM tree projects for which regulatory approval is currently sought

¹D: Development in contained laboratories; R: Release; T: Field testing; –: no information available.

²1 acre ~ 0.4 ha.

2.2.3.11 Adequacy of the existing framework (Q 3.3)

Respondents were asked about the regulatory framework in their country and whether the framework was adequate, from their point of view, to assess the benefits and risks related to their experiments. The evaluation of the response includes all respondents that are involved in genetic modification projects with trees (a total of 23 respondents). Three respondents
(13 percent) made no comments, five (22 percent) felt the framework was not adequate and 15 (65 percent) felt it was. Additional comments were offered on perceived shortcomings of the regulatory process and how improvements could be made. These included:

• process too slow, bureaucratic and costly

• exceptionally robust process, but too costly

• overcautious approach by regulatory agencies

• too many approvals required, often with the same assessment requirements, same information requested several times

• risk assessment is based on process – should be on product

• no consideration given to comparison with conventional methodology

• site safety concerns lead to requirement to keep location confidential

• policies need to be better publicized by regulatory agencies

• procedures need to be defined more clearly and made more efficient

• more risk assessment research required

• rules for contained research need to be relaxed

• applications need to be simplified for standard genes and promoters that have been assessed previously

• need to ensure that the process cannot be abused by opponents

2.2.3.12 Testing abroad (Q 3.4: Does the respondent have laboratories, field tests or commercial plantations abroad?)

None of the 23 respondents involved in genetic modification projects answered ‘yes’ to the question about whether they had research facilities abroad. Eighteen (78 percent) responded with ‘no’ and five (22 percent) did not respond at all. The responses were similar for field trials and commercial plantations. However, one respondent from Belgium reported that they have a field test in France. This respondent also confirmed that the regulatory process abroad was appropriate to assess the risk and benefit of the project.

2.2.3.13 Intellectual property regime (Q 4.1–4.3)

It is always difficult to obtain information related to intellectual property (IP). Much of this is confidential business information; however, respondents were encouraged to share information that they consider public. The responses from 23 respondents who use genetic modification in their projects were analysed. When asked whether they owned IP in relation to their production of GM trees, eight (35 percent) confirmed that they did, ten (44 percent) did not and five (22 percent) did not respond. The following list illustrates the broad areas in which IP is owned by these respondents:

• modulation of lignin biosynthesis

• transgene development

• vegetative multiplication

• testing of transgenes

• flowering control

• gene expression

• fibre development

• embryogenesis

• new varieties

• cell wall control

Nine of the 23 respondents (39 percent) confirmed that they have IP agreements with other institutions, eight (35 percent) did not and six (26 percent) did not respond. One respondent commented that since patent protection lasts for only 20 years and most GM trees grow slowly (many years to maturity), patent protection does not offer an advantage.

Eighteen respondents (78 percent) confirmed that they did not currently have IP arrangements with institutions in developing countries, five (22 percent) did not respond to this question. However, when additional comments were made, there appeared to be a preparedness to enter into IP arrangements where appropriate and required.

2.2.3.14 Future plans on use of genetic modification in trees (Q 5.1)

Here the organizations were asked for their plans on future use of genetic modification in trees. The analysis of responses included all respondents, since many organizations not using genetic modification at present made comments under this heading. The following lists represent the responses.

Prospects:

• 10-year timing horizon is envisaged to commercialization

• long term views (10–20 years) (20–30 years)

• “we have a restriction on research in British Columbia, a 10-year moratorium”

• research only, possibly field trials

• GM trees as research tools only, no commercialization anticipated

• concern that government regulation and public perception make tree genetic modification uneconomic

• GM trees as research tools only, no commercialization anticipated

• concerns related to unpredictability of expression

Developments:

• sterility to combat allergies

• assessment of environmental impacts

Potential products/uses:

• trees producing useful substances (for example pharmaceuticals)

• GM trees for site remediation (phytoremediation or bioremediation)

• use of genes for lignin reduction, increased yield and quality

• saving species (American chestnut, Castanea dentata, for example)

• assessing gene expression

• pest and disease resistance

• adaptive and fitness traits

• fibre quality

• resistance to dryness and high temperature

• candidate gene testing

• herbicide tolerance

It is difficult to quantify the responses in this category, but it is still interesting to note that many responses indicate an environmental or health benefit is expected from future use of GM trees.

2.2.3.15 Expected field release (Q 5.2: If and when a GM plantation is expected)

Only a few respondents predicted further field releases. One release is expected for 2004 and two further projects (potentially including many species) in around 10 years time. One respondent did not want to disclose information for reasons of confidentiality. Another respondent mentioned that no further tests would be conducted in the near future because of extremely high costs associated with field test applications (New Zealand). A respondent from Sweden felt that less than one percent of the Swedish forests would contain GM trees by 2050. However, this respondent also commented that in the long term GM trees will be planted more, because their characteristics can be manipulated more effectively.

Note that the terms ‘field test’ and ‘field release’ are not clearly defined. Some respondents regarded field tests as releases, others appeared to regard releases as full commercial plantations.

2.2.3.16 Benefits from GM trees (Q 5.3)

Respondents were asked about commercial, environmental, human health and other benefits of genetic modification of trees.

Several anticipated commercial benefits were mentioned by respondents using or not using genetic modification in tree projects. These are listed below in Table 2.2.8 (Figure 2.2.4) together with the number of times each expected benefit was cited.

Environmental benefits mentioned by respondents are listed in Table 2.2.9 (Figure 2.2.5). One respondent commented that 35 percent of the value of GM trees will lie in environmental benefits.

Human health benefits are probably more difficult to identify than commercial and environmental benefits. However, respondents made the comments listed in Table 2.2.10 (Figure 2.2.6).

Table 2.2.8. Commercial benefits anticipated from GM trees

Figure 2.2.4. Commercial benefits anticipated from GM trees (percentage of respondents citing benefit)

Table 2.2.9. Environmental benefits expected from GM trees

Figure 2.2.5. Environmental benefits expected from GM trees (percentage of respondents citing benefit)

Table 2.2.10. Human health benefits expected from GM trees

Benefit	Number of times cited
Pollen/allergy reduction	7
Environmental protection and restoration	5
Reduced environmental pollution	5
None/negligible/indirectly	3
Toxic site remediation	1
Pharmaceuticals	2
Recreation	1

Figure 2.2.6. Human health benefits expected from GM trees (percentage of respondents citing benefit)

Respondents were also asked to make any other comments with regard to benefits they would expect from GM trees in forestry. The following list reflects the responses:

• potential economic benefit to developing countries

• protection against insects and fungi

• significant acceleration of conventional breeding

• shortened breeding cycles

• focus by environmental groups on genetic modification has reduced their focus on clonal and conventional techniques

• knowledge and employment

• better forests, general improvements in forestry

• potentially decreasing global warming

2.2.3.17 Risks in GM tree use (Q 5.4)

Respondents were also asked to comment on potential risks in the use of GM trees in forestry.

Anticipated commercial risks are identified in Table 2.2.11 (Figure 2.2.7), human health risks in Table 2.2.12 (Figure 2.2.8), and environmental risks in Table 2.2.13 (Figure 2.2.9). Comments by respondents on other risks are also listed.

It is notable that the biggest threat to the commercial use of GM trees is seen to be associated with social rather than technical issues.

Table 2.2.11. Anticipated commercial risks of GM trees

Commercial risk	Number of times cited
Public resistance	9
High financial risk	5
Monopoly positions	3
Transgene instability	3
Lack of operational data	2
Plantation failure	2
Quality issues	2
Monocultures/biodiversity	2
Resistance development (insects)	1
Ecoterrorism	1
Low risk if managed properly	1

Figure 2.2.7. Anticipated commercial risks of GM trees (percentage of respondents citing risk)

Table 2.2.12. Anticipated human health risks of GM trees

Figure 2.2.8. Anticipated human health risks of GM trees (percentage of respondents citing risk)

Table 2.2.13. Anticipated environmental risks of GM trees

Environmental risk	Number of times cited
Limited/none	8
Gene escape	8
Plant escape, eco-disturbance	7
Impact on non-target species	2
Clonal failure	2
GM trees poorly adapted, so risk low	1
Toxic substances	1
Disturbance of food chains	1
Terrorism	1

Figure 2.2.9. Anticipated environmental risks of GM trees (percentage of respondents citing risk)

Further comments by respondents on ‘other risks’ (and in some cases their mitigation, which is dealt with in more detail in Table 2.2.14 [Figure 2.2.10]) are represented in the following list:

• GM forests may render natural forests valueless.

• Sterile trees may be developed to prevent or inhibit transgene spread.

• Government research priorities take funding and priorities and attention from other matters.

• Spread of competence in genetic modification increases risks connected with ‘bad will’ applications.

• Genetic modification will redirect resources from biological to legal matters.

• Legal and commercial matters reduce the potential for combining the best genes into one variety. This will delay release of varieties and promote the use of the same variety for a longer time, thus increasing the distance between technical progress and what is planted in the field.

• Genetic modification will encourage clonal forestry with a few clones that are not locally adapted (and used over a larger area than clones are currently).

• Radical actions against biotechnology as happened in the United Kingdom (in 1999), when extremists removed several crops of GM maize and a GM tree plantation.

• Given the invasive traits of some trees planted in commercial forests, there may be difficulties in controlling them if they are made genetically resistant to herbicides.

• Economic consequences are potentially high if resistance is broken by a new generation of parasites (whether resistance is given through genetic modification technology or not). This risk seems to be higher when (i) resistance is provided by a single gene, or a limited number of genes, (ii) for trees, compared with other plants, given the difference in growth times between a pest and its tree host, and (iii) when planted areas are significant (e.g. poplar and poplar leaf rust).

• Perhaps ecoterrorism.

The questionnaire also tried to gain some insights into how respondents felt risks could be addressed and how their R&D programme addressed the risks in particular. The results are compiled in Table 2.2.14 (Figure 2.2.10).

Table 2.2.14. How risks involved with tree genetic modification can be addressed

Figure 2.2.10. How risks involved with tree genetic modification can be addressed (percentage of respondents giving reply)

In a further question, respondents were asked to describe how their R&D programme addressed risk. The following list summarizes the responses:

• Research on flowering control.

• Study of pollen dispersal.

• Commercial trees will contain sterility genes when required.

• Use of D-serine resistance as a harmless selectable marker.

• By promoting biodiversity conservation as an issue to be dealt with within plantation forestry.

• Promoting plantation forestry as a tool for ecological restoration of native/natural forest types at risk.

• Study ecological interactions between GM tree material and herbivores (several insect species) and mycorrhizae.

• Collaboration with researchers working with societal and ethical questions related to GM plants.

• Cautious, step-by-step approach.

• “Significant R&D issue of the research programme of the Ministry of Consumer Protection; five research projects funded by the Ministry of Education and Research.”

• Safety assessment.

• Promoting forest tree genetic modification as an environmentally sustainable way to provide the world with timber, thereby creating the option to leave natural forests alone.

• “Even though we do not have an R&D programme, our objective is to serve as a facilitator for diverse groups to conduct appropriate research and to put the results into practice in an appropriate way.”

• No risk of genetic modification addressed since trees solely used for fundamental research.

At this point, the question was asked “what do you regard as the risks of not using genetic modification technology in commercial forestry?” Seven respondents felt that this would compromise the ability to reduce pressure on the world’s forests, three felt there were no risks at all, one regarded the risks as “very few” and another respondent did not foresee any economical risks. Further comments were made and they are summarized in the following list:

• Inability to compete at the highly intensive end of plantation production.

• Shortage of industrial wood during next century.

• Loss of a major industry in the United States to the tropics.

• Inability to save a species from extinction if it meets a serious threat (pest) to its survival.

• Depending on traits that are affected/modified, some environmental benefits are lost.

• Increases in pollution originating from traditional intensive forestry.

• Increased degradation of soil quality.

• “Intensive agriculture including forest plantation and the pulp and paper industries will always have a bad effect on the environment due to their intensive use of land resources and chemicals. Genetic engineering is the only way we can address these problems and reduce the damage.”

• Increased global warming and non-sustainable forestry will continue.

• Slowing down of traditional breeding programmes, while the demand for delivery of standardized products is always more pressing and urgent.

• “In a perspective of a few decades there is no risk, but in a longer time perspective we would abstain from the most powerful way of manipulating trees genetically.”

• “Genetic modification technology is just an extension of traditional tree breeding. To jettison the science would be a disservice to Mankind.”

2.2.3.18 The obstacles in the application of genetic modification to trees, and ways to overcome those obstacles (Q 5.5)

Respondents were asked about the obstacles they can see to applying genetic modification to forest trees.

Twenty-nine out of 49 (59 percent) of the respondents explained what they see as the major obstacles in using genetic modification in tree plantations, summarized in Table 2.2.15 (Figure 2.2.11).

Table 2.2.15. Obstacles to the use of genetic modification in trees. Most respondents identified more than one obstacle

Figure 2.2.11. Obstacles to the use of genetic modification in trees. Most respondents identified more than one obstacle (percentage of respondents citing obstacle)

When asked for suggestions on how to overcome these obstacles, respondents gave very complex answers. A total of 20 responses (41 percent of total) were received and 13 of them (65 percent) suggested better education of the public and communication with the public as a way to overcome obstacles. However, a number of other comments were made and they are listed below (in slightly edited form, to increase clarity):

• In the forestry sector, it would be wiser to tackle important traits, such as resistance/tolerance to drought/water stress, and to avoid gene transfer from distantly related organisms.

• Sterile/late flowering trees need to be developed.

• It might help if genetic modification were used with sterility genes, to limit genetic pollution.

• Implement a strategy for using an ‘antifitness-gene’ in combination with the ‘gene-of-interest’ to prevent the establishment of the transgenic line in natural ecosystems.

• Improve genetic modification technology in order to determine genetic transformation devoid of GUS (β-glucuronidase) and antibiotic resistant genes.

• Promote use of genetic modification technology only where there is a clear threat to the survival of a species (e.g. chestnut blight, Cryphonectria parasitica).

• “Can we increase the amount of conserved natural/old/wild forests/their biodiversity by producing the wood we need more effectively?”

• Avoid release of 'useless' GMOs, without obvious benefits to the final consumer/user (for example: resistance to herbicide).

• Close collaboration between pulp/paper industry and scientific research.

• Promote the possibility of certifying GM plantation products.

• Provide more R&D funds.

• Do not actively promote ‘commercial’ applications for GM forest trees over the next few decades (e.g. from FAO). Let the experiences from agriculture accumulate a little before looking for commercial applications. But do some field experiments to accumulate needed experience and knowledge and a better understanding of the consequences when commercial applications become interesting.

• “We need much more field-testing in order to gain experience, both of risks and benefits.”

2.2.3.19 Perception of the work done with GM trees (Q 6.1)

Respondents were asked whether public perception is an issue and whether they had experienced positive or negative attitudes towards their work. Out of the 23 working with tree genetic modification, 22 (96 percent) responded to this question. Also, three further respondents who did not work with tree genetic modification made comments. The following list of comments summarizes the responses. Note that the list does not contain all individual responses but it captures the main points made.

• NGOs are opposed to GMOs.

• “We have not had any negative comments directed at us because of our research, but then we are not carrying out field trials.”

• “Public perception is very important. Our institution is very active in science communication and has many different means of communication (website, leaflets, brochures, presentations, exhibition, school projects, etc.). GM trees are not a specific topic in this communication, but much information is provided on GM plants in general. With a transgenic poplar field trial, we experienced negative attitude from green activists. The trial was destroyed by these activists in July 1999.”

• Government agencies at local level strongly discourage non-research use of GM trees. At the national level, there is more support, but national agencies (Canadian Forest Service) have little power in provincial resource base decisions.

• “Public perception is important; forests are seen as our dear national property! Government agencies, I feel, do not understand our research work and its goals. The general public is interested and mostly relies on scientific risk assessments that regard ongoing research safe (we don't have commercial applications!). Then there is one small and very active activist group against GM trees, being also very skilful in getting a lot of publicity compared to their actual critical mass.”

• Positive, responsible attitude expressed by local authorities.

• “In Sweden the attitudes vary. Research was allowed. However, there was not enough evidence provided that GM trees are safe for the environment and human health. GM plants could not be planted in the field, only in greenhouses and incubators. Also there were newspapers reports of genetic modification of Norway spruce with a gene that confers freezing resistance. The gene was found in a deep-sea fish, so public opinion was that the trees would produce fish instead of cones and seeds. Outside the academic field, the perception of GM organisms is the same, as something terrible and evil, not for utilizing and even less for eating. This is the overriding opinion in my country.”

• Even though such activities are not under way in Romania, both forest authorities and the public have negative attitudes towards work on GM trees.

• “Yes, I do experience a negative attitude from the department, university and other parts of the establishment, because I say that I do not believe in fast application of genetic modification or the blessing of genetic patents as far as long-rotation forestry is concerned. As a result I get fewer research resources. But I feel that I should give my honest opinions on matters within forest genetics, when I feel there is a need to do so. From the ‘public’ (neighbours and so on) I meet a slightly negative attitude because they believe I am part of an ongoing transformation of the Swedish forest to GM forest, in spite of the fact that this initiative is a long way off.”

• The need for certification from the Forest Stewardship Council (FSC) has led Swedish forest companies to disassociate themselves from transgenic conifers even for research purposes (e.g. by refusing to allow trials of transgenic species on land supervised by the research organization).

• “We have experienced positive and negative attitudes towards our work. Mostly positive, except for members of Greenpeace.”

• “We are very concerned about public perception. A couple of years ago nearly 1 000 of our transgenic trees were vandalized by ‘eco’terrorists.”

2.2.3.20 Communication strategy (Q 6.2)

Respondents were asked whether their organization had a communication strategy to inform the public about developments in genetic modification. Nineteen out of 23 (83 percent) respondents working with GM trees made comments under this heading. Fourteen (74 percent) of them reported some form of direct communication, either as part of a corporate strategy or as individual communication strategies. Three (16 percent) respondents did not have any form of communication in the area of GM trees and two (11 percent) said that they relied on collaboration with outside agencies for their communication about GM trees.

2.2.3.21 Concluding comments (Q 7)

The opportunity was provided to make any additional comments that respondents wished to be considered in this study. The comments made here were mostly clarifications of what has been said earlier, and requests for copies of the study when available, plus requests for confidentiality.

2.2.4 FURTHER INFORMATION COLLECTED FROM PUBLIC SOURCES

To complement responses received to the questionnaire, and assess its overall representativeness, further information on field tests was collected from public sources in January 2004. They mainly included databases available on the Internet, often maintained by regulatory agencies (for detailed information and Internet addresses, see Appendix 3.8.1). Unfortunately, there is no database that covers all field tests worldwide and the individual databases have their own formats. The amount of data related to a specific trial varies to a great extent between the databases. Also, most often it cannot be determined from the database whether a specific trial has actually been established or whether its status is approved, but the trial not yet planted. However, in the following analysis all field tests listed in the databases are included and it is assumed that they have been planted or will be planted in the near future. Some databases did not appear to be up to date (they show field trial information up to a specific date which is sometimes years ago).

Both data sets referred to in this study (resulting from (i) the questionnaire and (ii) additional searches) were compared in detail to evaluate whether all trials specified in the questionnaire were actually covered in the additional analysis as well. Only two trials were not covered in the publicly available databasesTP.

All other trials that were reported in the questionnaire were also represented in the data set obtained independently from other resources. This indicates that the data in the questionnaire are very reliable. However, many (over 100) field trials found in other resources are not represented in the data set retrieved from the questionnaire.

2.2.4.1 Analysis of publicly available information

Numbers of field tests by country

In this analysis, a distinction was made between forest trees and a second category including both ornamentals and fruit trees (Table 2.2.16). Focusing on forest trees, the United States appear to have the majority of field trials active (103 trials reported in January 2004). Next is China with nine trials; two of them, a world-first, are commercial plantations. The countries in Europe, combined, have a total of 23 trials. For fruit trees and ornamentals, the United States is again the frontrunner with 47 trials, followed by Italy with eight trials.

It becomes evident from this analysis that the United States is under-represented in the responses to the questionnaire. It is of concern that the views of the major player(s) in forest biotechnology are not represented in this study, owing to a poor response from these organizations/companies. The views of researchers and organizations in the United States are poorly represented in the analysis of data from the questionnaire (especially from the private sector).

Table 2.2.16. Field trials of GM forest trees and GM ornamental/fruit trees recorded in publicly available databases, by country

GM genera/species involved

In the following analysis, forest trees were again separated from ornamental/fruit trees. It is noted that only a few forest trees are currently of significant interest for inclusion in forest biotechnology projects (Table 2.2.17). Populus is most often used in trials (82 trials), followed by Eucalyptus and Pinus (34 and 31 trials, respectively). In the fruit tree/ornamentals category, Malus is most often used in trials (33 trials), followed by Carica papaya and Prunus (18 and 13 trials, respectively).

Table 2.2.17. Field trials with GM forest trees and fruit trees/ornamentals

Genes/attributes involved

It is very difficult to extract comprehensive information on genes and traits from field-test databases (Table 2.2.18). It appears that those databases often contain some voluntary information that was provided by the organizations conducting tests, but this is not standardized and often one can only guess at the genes involved from the trait information given. Also, in many cases the genes are not specified (confidential business information). Further, it can be assumed that, at least at the time of the survey, many GMOs in field trials have an antibiotic resistance gene integrated into their genome (‘Reporter and marker genes’ in Table 2.2.18). This is important for selection of the transformed event and most researchers still use the nptII gene for this purpose. This gene confers resistance to the antibiotics kanamycin or geneticin. However, only 43 trials refer to this or a similar gene/trait. When assessing the additional traits of commercial importance to forestry, it is evident that herbicide resistance is the most important trait in forest trees (41 trials), followed by insect resistance (21 trials) and lignin modification (15 trials). In fruit trees/ornamentals the picture is significantly different with resistance against microorganisms (viruses, fungi, bacteria) appearing most important (35 trials), followed by insect resistance (11 trials).

Table 2.2.18. Genes and traits related to field tests

2.2.5 WHERE TO FROM HERE? THE FUTURE OF GENETIC MODIFICATION IN FORESTRY

The future of genetic modification in forestry appears difficult to predict. It is instructive, however, to assess the current situation in agricultural crops, where data from a decade of commercial application of genetic modification technology are now available (James 2004), along with ample reports on the environmental and economic sustainability of this technology. Most importantly, during this time the world has witnessed a continued and significant increase in GM crop acreage worldwide, with approximately 81 million hectares planted in 2004. Once plantations of GM crops were approved for a specified area or country, uptake by farmers has been swift, and more studies are forthcoming that substantiate the economic and ecological benefit of these crop plantations (FAO 2004).

Forestry is different in that rotation times are usually in the order of several decades (with the exception of short-rotation tree crops such as eucalypts). Nevertheless, where intensive plantation forestry can be practised, there are prospects of genetic modification eventually leading to field applications, provided objectives are well targeted and operational risks are managed appropriately. Studies have suggested high economic gain (Li et al. 2003), and comprehensive field trials with GM poplar with altered lignin composition have indicated that economic and environmental gains can be made, at least for producing pulpwood, with this technology (Pilate et al. 2002). As another example, field trials with GM Pinus radiata are beginning to yield data that demonstrate the continued correct expression of transgenes over 4–6 years and without selection for this expression (C. Walter, personal observation). The first commercial GM plantation, although relatively small in a plantation forestry sense (up to one million trees), is currently growing in China. GM Populus nigra trees have a Bacillus thuringiensis toxin gene integrated and the resistance of GM trees to insect attack has been demonstrated. More plantation projects using GM forest trees have been reported in China.

A number of issues that need special attention in the near future are detailed below, representing the authors’ personal analyses.

Biotechnologies including genetic modification have shown huge potential in the agricultural sector, to produce more food on less land, avert or mitigate certain undesirable environmental effects, and provide benefit to farmers, producers, consumers and the environment.

The Gene Revolution on-going in agriculture is fundamentally distinct from the Green Revolution of the 1960s and 1970sT in the sense that it is mainly driven by the private multinational sector, focuses on a very small number of crops and traits, and is protecting intellectual property in plant innovations (FAO 2004). Four countries (Argentina, Canada, China and the United States) and two traits (insect resistance and herbicide resistance) accounted for 99 percent of the global area planted with GM crops in 2003.

Molecular biology in forestry is now well advanced. Tree genomes and transcriptomes are being analysed and genetic transformation technology is available for many tree species, particularly those that are significant or potentially significant species for clonal plantation forestry.

Initial testing of GM trees in the laboratory, glasshouse and field has indicated that tree genetic modification can provide some tremendous advantages to growers and processors, while at the same time providing environmentally sustainable solutions for wood production.

Wood demand worldwide is on the increase, a trend that is likely to continue. The share of wood harvesting from natural forests will probably decrease, and be increasingly regulated. Forest biotechnology, in combination with plantation forestry, can provide options to protect natural forests from detrimental harvest while meeting the world’s increasing demand for wood.

New biotechnologies, in particular genetic modification, raise concerns. Admittedly, many questions remain unanswered for both agricultural crops and trees, and in particular those related to the impact of GM crops on the environment. Given that genetic modification in trees is already entering the commercial phase with GM Populus in China, it is very important that environmental risk assessment studies are conducted with protocols and methodologies agreed upon at a national level and an international level. It is also important that the results of such studies are made widely available.

2.2.6 ACKNOWLEDGEMENTS

The authors wish to acknowledge the contributions made by all respondents to the questionnaire. They have provided highly valuable information, which has made this study possible. Further, we thank Rowland Burdon, Mike Carson, Armin Wagner, Jens Find, Judy Griffith and John Smith for critically reading the report, making improvements and helping with graphics.

2.2.7 REFERENCES AND FURTHER READING

Beals, T.P. & Goldberg, R.B. 1997. A novel cell ablation strategy blocks tobacco anther dehiscence. Plant Cell, 9: 1527–1545.

Binns, A.N. & Thomashow, M.F. 1988. Cell biology of Agrobacterium infection and transformation of plants. Ann. Rev. Microbiol., 42: 575–606.

Bishop-Hurley, S.L., Zabkievicz, R.J., Grace, L., Gardner, R.C., Wagner, A. & Walter, C. 2001. Conifer genetic engineering: transgenic Pinus radiata (D Don) and Picea abies (Karst) plants are resistant to the herbicide Buster. Plant Cell Rep., 20: 235–243.

Burdon, R.D. 2001. Genetic aspects of risk – species diversification, genetic management and genetic engineering. N.Z. J. For., 44: 20–25

Burdon, R.D. & Walter, C. 2004. Exotic pines and eucalypts: perspectives on risks of transgenic plantations. In: S.H. Strauss& H.D. Bradshaw, eds. The bioengineered forest: challenges for science and society. Washington, DC, RFF (Resources for the Future) Press.

Carson, M., Carson, S. & Walter, C. 2004: The future of forest biotechnology – challenge paper. Proceedings, Forest Biotechnology Workshop, Global Biotechnology Forum, Concepcion, Chile, 2–5 March 2004.

Conner, A.J., Glare, T.R. & Nap, J.-P. 2003. The release of genetically modified crops into the environment. Plant J., 33: 19–46.

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

Dale, P.D. 1999. Public concerns over transgenic crops. Genome Res., 9: 1159–1162.

Dale, P.J., Clarke, B. & Fontes, M.G. 2002. Potential for the environmental impact of transgenic crops. Nature Biotechnol., 20: 567–574.

De la Cruz, I. & Davies, I. 2000. Horizontal gene transfer and the origin of species: lessons from bacteria. Trends Microbiol., 8: 128–133.

De Vries, J., Meier, P. & Wackernagel, W. 2001. The natural transformation of the soil bacteria Pseudomonas stutzeri and Acinetobacter sp. by transgenic DNA strictly depends on homologous sequences in the recipient cells. FEMS Microb. Lett., 195: 211–215.

Droege, W., Puehler, A. & Selbitschka, W. 1999. Horizontal gene transfer among bacteria in terrestrial and aquatic habitats as assessed by microcosm and field studies. Biol. Fertil. Soils, 29: 221–245.

Ellstrand, N.C. 2001. When transgenes wander, should we worry? Plant Physiol., 125: 1543–1545.

FAO. 2004. The state of food and agriculture 2003-2004. Agricultural biotechnology: meeting the needs of the poor? FAO Agriculture Series No. 35. Rome.

FAO. FAOSTAT interactive database (available at http://apps.fao.org/)

Fenning, T.M. & Gershenzon, J. 2002. Where will the wood come from? Plantation forests and the role of biotechnology. Trends Biotechnol., 20: 291–296.

Fischer, R., Buddle, I. & Hain, R. 1997. Stilbene synthase gene expression causes changes in flower colour and male sterility in tobacco. Plant J., 11: 489–498.

Fox, J. 2003. Resistance to Bt toxin surprisingly absent from pests. Nature Biotechnol., 21: 258–259.

Gianessi, L., Sivers, C.S., Sankula, S. & Carpenter, J.E. 2002. Plant biotechnology: Current and potential impact for improving pest management in US agriculture. An analysis of 40 case studies. Washington, DC, National Center for Food and Agricultural Policy. (available at www.ncfap.org/40CaseStudies.htm)

Gianessi, L., Sankula, S. & Reigner, N. 2003. Plant biotechnology: potential impact for improving pest management in European agriculture, a summary of nine case studies. , Washington DC, National Center for Food and Agricultural Policy, 15 pp. (available at www.ncfap.org/Europe.htm)

Grace, L.J., Charity, J.A., Gresham, B., Kay, N. & Walter, C. 2005. Insect-resistant transgenic Pinus radiata. Plant Cell Rep., 2005. (published online first)

Ho, M.-W., Ryan, A. & Cummins, J. 1999. Cauliflower mosaic viral promoter – a recipe for disaster? Microbiol. Ecol. Health Dis., 11: 194–197.

Hoefig, K.P., Moyle, R.L., Putterill, J. & Walter, C. 2003. Expression analysis of four Pinus radiata male cone promoters in the heterologous host Arabidopsis. Planta, 217: 858–867.

Hull, R., Covey, S.N. & Dale, P. 2000. Genetically modified plants and the 35S promoter: assessing the risk and enhancing the debate. Microbiol. Ecol. Health Dis., 12: 1–5.

Jain, R., Rivera, M.C. & Lake, J.A. 1999. Horizontal gene transfer among genomes: the complexity hypothesis. Proc. Natl. Acad. Sci. USA., 96: 3801–3806.

James, C. 2004. Preview: Global status of commercialized biotech/GM crops: 2004. ISAAA Briefs No. 32. ISAAA. Ithaca, NY, USA.

Krattiger, A.F. 1997. Insect resistance in crops: a case study of Bacillus thuringiensis (Bt) and its transfer to developing countries. ISAAA Briefs No. 2, 42 pp. Ithaca, NY, USA, International Institute for Applied Systems Analysis (IIASA).

Kube, P. & Carson, M. 2004. A review of risk factors associated with clonal forestry of conifers. In: C. Walter & M.J. Carson, eds. Plantation forest biotechnology for the 21PstP century. Kerala, India, Research Signpost.

Kumar, S. & Fladung, M. 2001. Gene stability in transgenic aspen (Populus). II. Molecular characterisation of variable expression of transgene in wild and hybrid aspen. Planta, 213: 731–740.

Levin, S.A. & Andreasen, V. 1999. Disease transmission dynamics and the evolution of antibiotic resistance in hospitals and communal settings. Proc. Natl. Acd. Sci. USA, 96: 800–801.

Levy, S.B., Marshall, B., Schluederberg, S. & Rouse, D. 1988. High frequency of antimicrobial resistance in human fecal flora. Antimicrob. Agents Chemother., 32: 1801–1806.

Li, L., Zhou, Y., Cheng, X., Sun, J., Marita, J.M., Ralph, J. & Chiang, V.L. 2003. Combinatorial modification of multiple lignin traits in trees through multigene co-transformation. Proc. Natl. Acad. Sci. USA, 100: 4939–4944.

Lindgren, K. & Lindgren, D. 1996. Germinability of Norway spruce and Scots pine pollen exposed to open air. Silva Fenn., 30: 3–9.

Lorenz, M.G. & Wackernagel, W. 1994. TBacterial gene transfer by natural genetic transformation in the environment. Microbiol. Rev., 58: 563–602.

Losey, J.E., Raynor, L.S. & Carter, M.E. 1999. Transgenic pollen harms monarch larvae. Nature (London), 399: 214.

Matzke, M.A. & Matzke, A.J.M. 1995. How and why do plants inactivate transgenes? Plant Physiol., 107: 679–685.

Meyer, P. 1995. Variation of transgene expression in plants. Euphytica, 85: 359–366.

Morel, J.-B. & Tepfer, M. 2000. Pour une evaluation scientifique des risques: le cas du promoteur 35S. Are there potential risks associated with use of the cauliflower mosaic virus 35S promoter in transgenic plants? Biofutur, 201: 32–35.

Nester, E.W., Thomashow, L.S., Metz, M. & Gordon, M. 2002. 100 years of Bacillus thuringiensis: a critical scientific assessment. American Academy of Microbiology, 18 pp. (also available at www.asm.org/Academy/index.asp?bid=2129)

Nielsen, K.M., Bones, A.M., Smalla, K. & van Elsas, J.D. 1998. Horizontal gene transfer from transgenic plants to terrestrial bacteria – a rare event? FEMS Microb. Rev., 22: 79–103.

Ochmann, H., Lawrence, J.G. & Groisman, E.A. 2000. Lateral gene transfer and the nature of bacterial innovation. Nature (London), 405: 299–304.

Owusu, R.A. 1999. Summary: GM technology in the forest sector, a scoping study for the WWF. 8 pp. (available at www.wwf-uk.org/filelibrary/pdf/gmsummary.pdf)

Phipps, R.H. & Park, J.R. 2002. Environmental benefits of genetically modified crops: Global and European perspectives on their ability to reduce pesticide use. J. Anim. Feed Sci., 11: 1–18.

Pilate, G., Guiney, E., Holt, K., Petit-Conil, M., Lapierre, C., Leple, 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 Biotechnol., 20(6): 558–560.

Pray, C.E., Huang, J., Hu, R. & Rozelle, S. 2002). Five years of Bt cotton in China the benefits continue. Plant J., 31: 423–430.

Punja, Z.K. 2001. GE of plants to enhance resistance to fungal pathogens – a review of progress and future prospects. Can. J. Plant Pathol., 23: 211–215.

Quist, D. & Chapela, I.H. 2001. Transgenic DNA introgressed into traditional maize landraces in Oaxaca, Mexico. Nature (London), 414: 541–543.

Rhymer, J.M. & Simberloff, D. 1996. Extinction by hybridization and introgression. Ann. Rev. Ecol. Syst., 27: 87–109.

Sedjo, R. 2004. Potential for biotechnology applications in plantation forestry. In: C. Walter & M.J. Carson, eds. Plantation forest biotechnology for the 21st century. Kerala, India, Research Signpost.

Shin, D.I., Podila, G.K., Hunag, Y. & Karnosky, D.F. 1994. Transgenic larch expressing genes for herbicide and insect resistance. Can. J. For. Res., 24: 2059–2067.

Snow, A. 2002. Transgenic crops – why gene flow matters. Nature Biotechnol., 20: 542.

South, D.B. 1999. How can we feign sustainability with an increasing population? New For., 17: 193–212.

Stewart, C.N., Richards, H.A. & Halfhill, M.D. 2000. Transgenic plants and biosafety: Science, misconceptions and public perceptions. BioTechniques, 29: 832–843.

Strauss, S.H., Rottmann, W. H., Brunner, A.M. & Sheppard, L.A. 1995. GE of reproductive sterility in forest trees. Mol. Breed., 1: 5–26.

Tang, W. & Tian, Y. 2003. Transgenic loblolly pine (Pinus taeda L.) plants expressing a modified delta-endotoxin gene from Bacillus thuringiensis with enhanced resistance to Dendrolimus punctatus Walker and Crypyothelea formosicola Staud. J. Exp. Bot., 54: 835–844.

Tepfer, D., Garcia-Gonzales, R., Mansouri, H., Seruga, M., Message, B., Leach, F. & Mirna Curkovic, P. 2003. Homology-dependent DNA transfer from plants to a soil bacterium under laboratory conditions: implications in evolution and horizontal gene transfer. Transgen. Res., 12: 425–437.

Thompson Campbell, F.T. 2000. Genetically engineered trees: questions without answers. Washington, DC, American Lands Alliance (available at www.americanlands.org/forestweb/getrees.htm)

Van den Belt, H. 2003. Debating the precautionary principle: “Guilty until proven innocent” or “ Innocent until proven guilty”? Plant Physiol., 132: 1122–1126.

Walter, C. 2004. Review: genetic engineering in conifer forestry: Technical and social considerations. In Vitro Cell Dev. Biol.-Plant., 40: 434–441.

Walter, C. & Fenning, T. 2004. Deployment of genetically-engineered trees in plantation forestry – an issue of concern? The science and politics of genetically modified tree plantations. In: Walter, C. & Carson, M.J., eds. Plantation forest biotechnology for the 21Pst century. Kerala, India, Research Signpost.

Whiteman, A. & Brown, C. 1999. The potential role of forest plantations in meeting future demands for industrial wood products. Int. For. Rev., 1(3): 143–152.

Su, X.H., Zhang, B.Y., Huang, Q.J., Huang, L.J. & Zhang, X.H. 2003. Advances in tree genetic engineering in China. Paper submitted to the XII World Forestry Congress, Quebec, Canada. (www.fao.org/DOCREP/ARTICLE/WFC/XII/0280-B2.HTM)

Yasueda H., Yui, Y., Shimizu, T. & Shida, T. 1983. Isolation and partial characterization of the major allergen from Japanese cedar (Cryptomeria japonica) pollen. J. Allergy Clin. Immunol., 71: 777–786.

Zhu, S., Tomberlin, D. & Buongiorno, J. 1999. Global forest products consumption, production, trade and prices: global forest products model projections to 2010. FAO Global Forest Product Outlook Study Working Paper No: GFPOS/WP/01. Rome

2.2.8 ANNEXES

2.2.8.1 Resources

Field test approval authorities (as advised by questionnaire responses, plus additional resources).

Internet addresses of organizations publishing information on the Internet (compiled from responses to questionnaires.

Country	Organization	Internet address
Australia	CSIRO Forestry and Forest Products, PO Box Kingston, ACT 2604, Australia	www.ffp.csiro.au/tigr/molecular/molbiology.html
Belgium	Department of Plant Systems Biology, Vlaams Instituut voor Biotechnologie (VIB), Rijvisschestraat 120, 9052 Zwijnaarde, Belgium	www.psb.rug.ac.be/; Follow - Research: Research divisions: Molecular Genetics: Tree Biotechnology to find research details
Canada	Atlantic Forestry Centre, PO Box 4000, Fredericton, NB, Canada E3B5P7	www.nrcan.gc.ca
Finland	Punkaharju Research Station, Finlandiante 18, FIN-58750, Punkaharju, Finland	www.metla.fi/tutumus/index-en.htm; search for "biotechnology' or Project 3198 under the topic ‘Forest Genetics and Tree Breeding’
France	Unité Amélioration Génétique et Physiologie Forestières, INRA-Orléans, Avenue de la Pomme de Pin, BP20619 Ardon, 45166, Olivet Cedex, France	www.inra.fr/
	AFOCEL Biotech, Domaine de l'Etancon, F-77370 Nangis, France	www.afocel.fr/
Germany	Institute for Forest Genetics and Forest Tree Breeding, Federal Research Centre for Forestry and Forest Products, Sieker Landstr. 2, D-22927 Grossgansdorf, Germany	www.rrz.uni-hamburg.de/GeneTree/
Japan	Forestry and Forest Products Research Institute, Matsunosato 1, Tsukuba, Ibaraki-ken, Japan 305-8687	http://ss.ffpri.go.jp/e_version/majorprograms/mjprograms_j
New Zealand	The Horticulture and Food Research Institute of New Zealand Ltd, Private Bag 92 169, Auckland, New Zealand	www.hortresearch.co.nz
	New Zealand Forest Research Institute Ltd, Sala Street, Rotorua, New Zealand	www.forestresearch.com
Sweden	Department of Plant Biology and Forest Genetics, Swedish University of Agricultural Sciences, Box 7080, S-750 07 Uppsala, Sweden	www.genfys.slu.se
USA	Institute of Forest Biotechnology, PO Box 13399, Research Triangle Park, NC 27709-339, USA	www.forestbiotech.org
	Tree Genetic Engineering Research Cooperative, 348 Richardson Hall, Oregon State University, Corvallis, OR 97331-5752, USA	www.fsl.orst.edu/tgerc/index.htm

²⁸ Contact: [email protected].

²⁹ Contact: [email protected].

³⁰ IPGRI’s European Forest Genetic Resources Programme.

³¹ For examples, see the websites of regulating authorities in Appendix 3.8.1.

³² Numbering refers to the numbers of the questions in the questionnaire.

³³ Note that percentages add to more than 100 because most respondents work on more than one species.

³⁴ These data are probably a vast underestimate of the actual number of trees installed in field trials.

³⁵ The cry1Ac gene.

³⁶ Cry1Ac codes for a protein that is toxic to a specific class of insects.

³⁷ Another gene of which the product is toxic to insects.

³⁸ Non-governmental organizations.