3.1 BACKGROUND DOCUMENT
3.2 SUMMARY DOCUMENT
HOW APPROPRIATE ARE CURRENTLY AVAILABLE BIOTECHNOLOGIES FOR THE FORESTRY SECTOR IN DEVELOPING COUNTRIES?
Plant biotechnology is a field of scientific research in which rapid advances have been made in recent years and which appears to have much potential for further development. Numerous opportunities for using biotechnology in plant breeding have been identified, some of which might be appropriate for the improvement of crops in developing countries, as discussed in the crop sector conference (Chapter 2). In this conference the focus is on forest trees and currently available biotechnologies and their application in the forestry sector are discussed with reference to their potential use in developing countries today. Please note that for the purposes of this conference, the term forestry sector specifically excludes fruit orchards.
Most forest tree species are characterized by inherently high levels of variation and extensive natural ranges. This high level of genetic variation needs to be maintained to ensure present-day and future adaptability to changing environmental conditions. It is also needed to maintain options and potential for improvement to meet changing end-use requirements. Forests provide a wide range of goods and services such as timber, fibre, fuelwood, food, fodder, gum, resins, medicines, pharmaceutical products and environmental stabilization. Similar goods and services are often provided by a wide range of genera and tree species. Despite the availability of a large number of forest tree species, less that 500 have been systematically tested for their present-day utility for human beings and less than 40 species are included in intensive selection and breeding programmes.
Selection in breeding populations with a broad genetic base is the most common approach to forest tree improvement. Although demand for wood is the driving force in the development of large-scale forest plantations, several selection and breeding programmes aim at enhancing other goods and environmental services provided by forest trees and shrubs.
Since most forest tree species are characterized by long generation intervals and a generally long juvenile phase before flowering, much time is needed before assessment of important traits can be carried out. For example, if wood quality is of interest in breeding for timber or fuelwood, selection can only be carried out after trees have reached a certain size which, in some cases, can require decades. The above factors are limitations to rapid improvement and only a maximum of three or four generations of breeding have been completed in a few forest tree species to date (Eucalyptus grandis and some pine species).
3.1.2 Description of biotechnologies in the forestry sector
This section provides a summary of recently developed biotechnologies that could be used, or more widely used, for forest trees in developing countries today. Additional technical details may be found at http://www.fao.org/forestry/FOR/FORM/FOGENRES/genresbu/125/125e/arte11.stm
188.8.131.52 Biotechnologies based on molecular markers
Reliable information on the distribution of genetic variation is a prerequisite for sound selection, breeding and conservation programmes in forest trees. Genetic variation of a species or population can be assessed by measuring morphological and quantitative characters in the field or by studying molecular markers in the laboratory. A combination of the two methods is required for reliable results.
Molecular markers can be used for:
a) Quantification of genetic diversity
The use of molecular markers for the determination of the extent of variation at the genetic level, within and between populations, is of value in guiding genetic conservation activities, which are aimed at maintaining genetic diversity with respect to traits of both known and unknown importance, and in the development of breeding populations for specific end uses.
It should be noted that studies on genetic diversity based on molecular markers must be interpreted with caution, due to frequently low correlations with patterns of variation for adaptive traits, which are of major importance in forestry.
b) Genotype verification
Molecular markers have been widely used for identification of genotypes and applied in taxonomic studies, biological studies and genetic fingerprinting. Good taxonomy is fundamental to conservation and tree improvement programmes and to programmes involving hybridization between species. The use of molecular markers has revolutionized studies of mating systems, pollen movement and seed dispersal. Results of such biological studies are of considerable practical significance to advanced tree improvement programmes, specifically in population sampling, design and management of seed orchards (i.e. orchards consisting of clones or seedlings from selected trees and cultured for early and abundant production of seeds for reforestation), estimation of pollen contamination and development of controlled pollination methods. Germplasm identification, through genetic fingerprinting, has been used in advanced breeding programmes which rely on controlled crosses or in which the correct identification of clones for large-scale propagation programmes is essential.
c) Gene mapping and MAS
Genetic linkage maps can be used to locate genes affecting quantitative traits of economic importance. Quantitative traits, such as wood yield, wood quality or pulp yield, are usually controlled by many genes, termed QTLs. By using molecular markers closely linked to, or located within, one or more QTL, information at the DNA-level can be used for early selection. The potential benefits of MAS are greatest for traits that are difficult, time-consuming or expensive to measure (for example, wood quality traits or pulp yield). Mapping and MAS tend to be used only in species of high economic value and have most potential in clonal breeding programmes, where additional genetic gains can be rapidly multiplied.
184.108.40.206 Biotechnologies based on vegetative propagation
Strategies supporting large-scale utilization of genetic material with a narrow genetic base must be appropriately integrated into tree improvement programmes. Vegetative propagation within such programmes allows for a fast release of new materials and for appropriate matching of clones to different local conditions. It also allows continued cultivation of given clones and to efficiently change the mixture of clones used in a given programme. Vegetative propagation also supports other currently available biotechnologies (in vitro storage and cryopreservation; in vitro selection).
a) In vitro storage and cryopreservation
In vitro storage refers to the storage of germplasm in aseptic culture under laboratory conditions, while cryopreservation refers to the storage of cells, tissues, seeds, etc. at temperatures of liquid nitrogen (-196ºC). The two techniques do not seem to be widely used in genetic conservation activities for forest trees, but they may serve as back-up strategies for species with seed storage problems.
b) In vitro selection
In vitro selection refers to the selection of germplasm based on test results using tissue culture under laboratory conditions. Many recent publications for crop plants have reported useful correlations between in vitro responses and the expression of desirable field traits, most commonly disease resistance. Positive results are available also for tolerance to herbicides, metals, salt and low temperatures. For the selection criteria of major general importance in forest trees (in particular vigour, stem form and wood quality), poor correlations with field responses will limit the usefulness of in vitro selection. However, in vitro selection may be of possible interest in forestry programmes for screening disease resistance and tolerance to salt, frost and drought.
For crop and horticultural species, micropropagation (in vitro vegetative propagation of plants) is now the basis of a large commercial industry involving hundreds of laboratories around the world. Successful protocols now exist for a large number of forest tree species and the number of species for which successful use of somatic embryogenesis has been reported is increasing (somatic embryogenesis is a step in micropropagation where somatic cells are differentiated into somatic embryos). So, in the future, it is likely that micropropagation in the forestry sector will become commercially more important. Compared to vegetative propagation through cuttings, the higher multiplication rates available through micropropagation seem to offer a quicker capture of genetic gains obtained in clonal forestry strategies.
One major factor impeding early application of micropropagation in many large-scale forest plantation programmes, is that breeding and selection of desired clones are not sufficiently advanced for clonal forestry to be contemplated. Current high costs will also be an impediment to the direct use of micropropagation in many programmes. Technologies resembling those used commercially in horticulture are most likely to be affordable for a limited number of high-value forest tree species, particularly those for which propagation by cuttings is difficult. Micropropagation is unlikely to be used for the production of planting stock of non-industrial forest tree species.
220.127.116.11 Genetic modification of forest trees
GMOs are defined as organisms that have been modified by the application of recombinant DNA technology (where DNA from one organism is transferred to another organism). The term transgenic trees is also used for GM trees, where a foreign gene (a transgene) is incorporated into the tree genome.
One of the first reported trials with GM forest trees was initiated in Belgium in 1988 using poplars. A study carried out in 1999 indicated that, since then, there have been more than 100 reported trials, involving at least 24 tree species - most of which are timber-producing species. The majority of the field trials were carried out in the USA and Canada. Whereas it is estimated that roughly 40 million hectares of transgenic agricultural crops were grown commercially in 1999 (ISAAA figures), there is no reported commercial-scale production of transgenic trees. Information on field trials of GM trees has been published by the OECD [www.olis.oecd.org/biotrack.nsf] and the World Wide Fund for Nature (1999) [www.wwf-uk.org/news/n_0000000172.asp].
Traits for which genetic modification can realistically be contemplated in the near future include insect and virus resistance, herbicide tolerance and lignin content. However, insertion of any gene into a tree species with expected functional results will be a substantial undertaking and insertion of enough genes to confer e.g. long-term insect resistance in a perennial species even more so. Virus and insect resistance, in particular, are of major significance for crop plants. By contrast, these traits are not the most important in breeding programmes of forest tree species (poplars being an exception). Reduction of lignin is a valuable objective for species producing pulp for the paper industry; work on this aspect is underway in aspen.
A major technical factor limiting the application of genetic modification to forest trees, is the current low level of knowledge regarding the molecular control of traits which are of most interest, notably those relating to growth and stem and wood quality. Genetic modification of these traits remains a distant prospect. Investments in genetic modification technologies should be weighed against the possibilities of exploiting the large amounts of genetic variation, which are generally untapped, available within any single species in nature.
Biosafety aspects of GM trees need careful consideration because of the long generation time of trees, their important role in ecosystem functioning and the potential for long distance dispersal of pollen and seed.
3.1.3 Forestry in developing countries
Forests cover approximately 30 percent of the worlds total land area [Data from 2000, see www.fao.org/forestry/fo/sofo/sofo-e.stm]. They are the source of vital commodities, including raw materials and food and are essential for maintaining agricultural productivity and the environmental well-being of the planet as a whole. They protect soil and water and buffer the effects of wind and rain, thus helping to decrease soil erosion and they are an important sink for carbon dioxide. Forests are also among the most important repositories of biological diversity.
Roughly 500 million rural people live in, or close to, forests. Most communities use a variety of forest products, particularly those in developing countries. Plant stems, tubers and fruits provide additional food during hungry seasons or when crops fail; wild animals are harvested for meat and hides; and the forests provide fuelwood, fodder for livestock, medicines and other products and services.
The most important trend in forestry in developing countries is the progressive reduction in the area of forests due to changes in land use. Another important trend, evident at a global level, is increasing forest degradation through unmanaged use. When forests are degraded, their productive functions and their capacity as regulators of the environment are reduced, increasing flood and erosion hazards, reducing soil fertility and contributing to the loss of forest products and overall loss of biological diversity.
While forests are being lost, there is growing demand both for environmental services and for wood and wood products which they provide. A forecast by FAO predicts that wood demand is expected to increase by 25 percent from 1996 to 2010. This demand will, increasingly, have to be met by forest plantations, and with decreasing land areas available for forestry, plantation methods will have to be increasingly intensive. This will necessitate better tree improvement programmes in which biotechnology may play a role.
3.1.4 Certain factors that should be considered in the discussion
The key question in this e-mail conference is how appropriate each of the different biotechnologies may be for the forestry sector in developing countries today.
The question of appropriateness should consider the following elements:
As was the case for the e-mail conference on crops (Chapter 2), this conference asked a similar question, i.e. how appropriate are currently available biotechnologies for the forestry sector in developing countries? Again, the three major areas of discussion revolved around biotechnology based on the use and development of a) molecular genetic markers; b) micropropagation; and c) GM trees. However, the technology of genetic modification was by far the primary topic of discussion.
Thirty-two submissions were received during the e-mail conference, compared to 138 in the crop conference, but the 32 messages covered a wide range of ideas related to the three major areas. Comments ranged from general observations to very detailed suppositions. Important points were made several times and these formed the basis for themes that emerged.
Section 3.2.2 of this document attempts to summarize these themes. Section 3.2.3 provides additional points that did not fall logically into the general themes, but were important points to consider. Specific references to messages posted, giving the participants surname and the date posted (day/month of the year 2000), are included. The messages can be viewed at www.fao.org/biotech/logs/c2logs.htm. Section 3.2.4 gives the names and country of the people that sent referenced messages.
3.2.2 Major themes and factors of importance for the application of biotechnology
18.104.22.168 All biotechnologies need to be considered within the framework of a larger genetic resource management programme
This point was made several times, i.e. modern biotechnology should only be realistically developed for species which already have a substantial infrastructure in basic plantation technology, e.g. in seed collection, nursery techniques, silviculture and in tree breeding and related research.
Serrano (9/5) indicated that while research is underway in somatic embryogenesis in pine in Mexico, the largest problem is that of basic forest management practices (e.g. appropriate harvesting systems). This may point out a fundamental dilemma for developing countries with respect to investments in biotechnology. If there are more basic forest management issues to be solved, then should investments be made in technologies that may never be applied? Burdon (20/6) added to this by saying, in the short- to medium-term, the development of biotechnology is likely to make much increased demands of the breeding infrastructure.
Southerton (19/6) emphasized this point again by saying that there is a danger in rushing to use the latest technologies when more basic approaches, such as provenance testing (i.e. seed source) and selection of appropriate plantation species, would provide a much larger payoff. Ashton (13/6) suggested that the discussion may be premature for forestry at this time (e.g. it is not yet simple to transfer multiple-gene constructs to a recipient genome), so developing countries need to focus more on recreating the full local diversity of forest ecosystems rather than genetically engineering some unstable, unpredictable exotic import.
Several contributors appropriately pointed out that many GM transformed clones would have to be developed for use at any one time, and most would be screened out due to poor performance or poor stability. Smith (15/6) suggested that if there are additional concerns about the use of GM trees (over and above simple clonal forestry), there could be requirements for increasing the rigour and length of time for field testing protocols. This could put the utility of technology in an even more cost prohibitive situation. DiFazio (7/6), Strauss (7/6) and Smith (15/6) all agreed that deployment and monitoring schemes that address appropriate genetic considerations for safety and productivity of GM tree plantations have to be developed and implemented. Furthermore, as suggested by Strauss (7/6), and supported by Hong (8/6), the assessment of risk could be responsibly monitored if there are step by step requirements laid out: the same requirements that should apply to any good silviculture or breeding programme.
Even in more developed tree breeding programmes, a push to develop advanced techniques such as markers for QTL selection could increase demands on tree breeding programmes (e.g. larger progeny tests required). Burdon (20/6) summarized this theme quite appropriately by saying, the application of new biotechnology will need to stand as an enhancement of classical breeding, rather than as a substitute for it.
22.214.171.124 Long rotation ages for most forest tree species
Lindgren (4/5) made several observations related to the use of new biotechnology and the long generation time of tree species relative to crop species. He noted that, first, many developing countries are in warmer climates and many of the species used by them may have relatively short rotation ages (rotation age is the age at which trees are harvested). GM trees with short rotation ages would also be more reliable, with respect to expression of the trait (i.e. testing ages may more closely correspond to harvesting ages, so there would be greater confidence in trait expression). For long-rotation species, there would be many doubts, because testing would probably not be able to cover the full rotation (which is particularly important if the trait is required for the full lifetime of the tree). Second, some of the end-product objectives of GM trees, e.g. special pulping attributes, are likely to be more relevant for short rotation species. Third, even for some of the major commercially important pine species (with rotation ages typically over 20 years), investments in new biotechnologies may not be profitable. However, he proposed that it could be appropriate for those species that will be tested and harvested within a roughly 10-year time frame (Moderators note: we are assuming this approximation would need to be based on some investment calculations).
Strauss (10/5, message 4) stated that GM trees will be limited to the common short-rotation forest tree species in intensively grown plantations in the developing world (e.g. Eucalyptus) and intensively managed species (e.g. poplars and some pines) in the developed world. Later (7/6), he re-iterated the point that GM trees will only be used commercially after a number of years of testing on many sites. During this process the vast majority of transgenic lines are discarded......only those that are most stable and perform well are considered for commercial use. Lindgren (14/6), supported by Southerton (19/6), pointed out that there would be a tendency to use fewer clones, so it may be best to see how trends develop for clonal forestry programmes (e.g. in Eucalyptus) around the world. Again, this stresses the need, as mentioned earlier, for developing genetic diversity and deployment guidelines. Strauss (7/6) stressed the basic fact that there are also substantial physical limitations to establishing large areas of forest tree plantations on a scale and time frame similar to that of crop species.
In summary, lengthy research and developmental periods will be required to develop and deploy GM trees. Therefore, it is likely that there will be a relatively long time period for foresters, relative to crop geneticists and agriculturists, to monitor and correct trends and policies in the use of GM trees, prior to large-scale use across plantation estates.
126.96.36.199 Technology being appropriate or inappropriate for developing countries
There was a clear consensus that many factors need to be considered in deciding whether or not any biotechnology is appropriate in forestry (i.e. biological, economical, and political restraints and opportunities). Therefore, it was not easy to say that modern biotechnology is either appropriate or not appropriate for developing countries.
As mentioned above, Lindgren (4/5) argued that although developing countries may not generally have advanced infrastructure and modern laboratories, they often have better growing conditions for trees (shorter rotations) than temperate/boreal developed countries. Strauss (7/6) noted this is particularly relevant for Eucalyptus in some developing countries, in which well-developed plantation forestry systems are already in place.
Keeping local options open was brought up a few times. As pointed out by Strauss (10/5, message 3), why do we seek some kind of global consensus about use of genetically engineered plants and trees?. He added, all practitioners know, the only place [GM trees] will find use, for the foreseeable future, is in intensively managed plantations - whether they be industry or community owned. Fenning (14/6) further supported this view by stating that people should be left free to choose the most appropriate solution to local needs in future.
Another view of the issue was that if the appropriate technology exists for a given situation, it would be negligent not to apply the tools available (Fenning, 9/6). For example, some modern tissue culture techniques may be suitable for special situations; such as the conservation and management of Prunus africana (Smith (11/5)), which has been used for medicinal purposes and may require special attention to ensure sustainability of the resource.
This does, however, raise the general concern of whether developing countries have the means and resources to appropriately assess or manage the risks, compared with more developed countries. This was to some degree raised by Johnston (11/5), who stated that the burden of proof for risk assessment should lie with the proponents of the technology. Smith (29/6) also pointed out that technologies might have additional hidden costs in the future and not just environmental risks. For instance, products from the early attempts at tissue culture showed that physiological ageing was present that could reduce stem volume growth in trees produced by tissue culture. (This may not be detected in the testing phase). An additional point is that even with the use of conventional technology (e.g. fast growing plantation management), the characteristics of the wood may change and require research and development in processing technology (e.g. special drying/sawing technology). These issues may be risky for developing countries that may not be able to bear the additional costs of research and development for a changed product.
188.8.131.52 Increased public awareness and societal concerns regarding the threats and benefits of biotechnology
Nine of the 32 e-mail submissions touched upon this general concern. A quote from Strauss, Raffa and List (26/5) is quite appropriate to sum up the general concerns of this theme:
The challenges to ethical uses of GM trees in forestry reside not in the process by which they are created, but rather in how their new characteristics and use will affect the environment and society. Substantial benefits have been documented in laboratory and field experiments. However, there are reasonable ecological and social concerns based on precedents from other kinds of agricultural technology. The key problems are deciding when our knowledge base is adequate, when there has been sufficient public discussion, and when there is adequate social consensus that the net effects for proposed uses are positive. If the process of public evaluation is scientifically sound and democratically rigorous, it should be possible to enjoy a continuing flow of new products from this rapidly maturing technology for the benefit of forestry in coming decades. If it is not, the technology may remain on the shelf in spite of its technical merits.Johnston (11/5) agreed that decisions regarding biotechnology should be made on local needs, economics and environmental considerations and that all the risks and alternatives must be discussed alongside the possible benefits. Overall, there was a large consensus that there is a much greater need now for public information and awareness of these technologies, before they should or will be used. Although most, if not all, GM trees will be used in high investment plantations, there are complex ecological questions that still must be carefully analysed.
184.108.40.206 Ownership and sharing of germplasm, techniques and financial arrangements with developing countries
Compared to the crop conference, there was rather limited discussion on the problem of moving new technologies (e.g. genetic modification) to developing countries. Perhaps it is not as clear in forestry, with respect to where specific genetic modification technologies would be useful, as no GM trees have yet been commercially released.
In some developing countries, ownership of land, forests and trees is not clear. This was specifically raised by Fenning (19/5) who said that it may not be clear who owns the forests or trees in developing countries where this technology could be applied. This creates a fundamental problem of guarantees on who will actually reap the benefits from any specific investment in these situations.
Fenning later (14/6) suggested (a point also made in the crop conference), that there is a need for innovative ways to provide access to the appropriate biotechnology for local programmes in developing countries. There were, however, no real proposals or examples in the e-mail conference where this was examined. Smith (13/6) pointed out that patent lives of around 20 years could provide substantial protection for certain types of biotechnologies. However, this may not be directly applicable to forestry, as trees planted 20 years from now with the patented technology, or those developed now but which are not harvested till later (after more than 20 years), may not be subject to such patent protection (or financial obligations or previous arrangements to the patent holders). In the short-term, patent or ownership restrictions could have immediate effects on investment incentives, particularly if there are large upfront costs associated with purchasing rights to use various products or techniques of biotechnology.
Burdon (19/5), considering political and institutional aspects of biotechnology, wrote the following that summarizes the issue quite nicely:
Much may depend on the agencies involved. If large foreign investors are involved, they can in principle put in place a well-balanced technological base, whereby the biotechnology is properly coupled with complementary, field-based programmes in which there is a proper infrastructure of genetic management. However, for such an organization, the operation in a single developing country may be a small part of a global risk spread, in contrast to the risk exposure for the individual country and especially the local community(ies). In this situation there will also be Intellectual Property issues, while the regulatory mechanisms for risk management (which is not straightforward anywhere) are likely to be weak.3.2.3 Additional points of relevance to the use of biotechnology in forestry
3.2.4 Name and country of participants with referenced messages
Burdon, Rowland. New Zealand
DiFazio, Steve. United States
Fenning, Trevor. Germany
Hong, L.T. Malaysia
Immonen, Sirkka. Italy
Johnston, Sam. United States
Lindgren, Dag. Sweden
Serrano, Carlos Ramirez. Mexico
Smith, Dale. New Zealand
Southerton, Simon. Australia
Strauss, Steven. United States
Strauss, Steven; Raffa, Kenneth and List, Peter. All from United States