J. D. MATTHEWS
J. D. MATTHEWS is Professor of Forestry, University of Aberdeen, Old Aberdeen, Scotland, and leader of Section 22 of the International Union of Forest Research Organizations. He served as general rapporteur to the Consultation and has edited this final report. He acknowledges with thanks the assistance of his colleagues, J. S. Murray and E. Innes, the co-operation of the authors of the papers which follow, and the advice of A. Metro of the Forestry and Forest Products Division of FAO.
With the government of Sweden acting as host, and with the support of the International Union of Forest Research Organizations, a World Consultation on Forest Genetics and Tree Improvement was held in Sweden from 20 to 30 August 1963. This important meeting was organized by FAO in response to a recommendation by the Fifth World Forestry Congress in Seattle, in 1960. The Congress recognized the need for action to use forest tree improvement to strengthen afforestation and reforestation programs and to increase the yield of the world's forests. It therefore recommended that a worldwide technical conference be organized by FAO to co-ordinate and promote the development of tree improvement techniques, the mass production of improved planting stock, and the incorporation of these techniques and plants into afforestation and reforestation programs in a scientific and economic manner.
Detailed preparations for the meeting began in May 1962 and the agenda was worked out at a meeting which was held at FAO headquarters and attended by Å. Gustafsson representing Sweden, A. de Philippis, P. Bouvarel and J. D. Matthews representing IUFRO, and A. Metro, M. Andersen and O. Fugalli of the FAO Forestry and Forest Products Division. A series of leading and supporting papers were solicited from research workers active in the chosen aspects of forest genetics and tree improvement. The leading papers were written in the form of draft chapters for the final report of the Consultation and were later revised and rewritten by their authors to incorporate the results of the Stockholm discussions, which were summarized by the drafting teams, and the salient points of the supporting papers.
A total of 119 solicited and voluntary papers were written for the Consultation. Those listed in Appendix D (Sections 1-8) of this final report have been published in full as the proceedings of the Consultation.1 Mention must also be made of the valuable background papers produced for the three study tours by the Swedish Organizing Committee and their colleagues. These papers are tasted in Appendix D (Section 9) and the texts of three of them appear as Appendixes A, B and C of this final report.
1 Proceedings of the World Consultation on Forest Genetics and Tree Improvement, FAO, 1964.
The conference proper was preceded by three short study tours. The first went to northern Sweden with Thorsten Andrén as Chairman and Fritz Bergman as technical leader. It was attended by 50 participants, who saw seed orchards, progeny testing, the experiment station of the Swedish Forest Tree Breeding Association at Sundmo, provenance tests and related sample plots. The second study tour started from Karlstad in central Sweden with Sven G. Ekman as Chairman, Erik Edlund as technical leader and Tore Arnborg as technical consultant to the 40 participants. The program included visits to the experiment station of the Swedish Forest Tree Breeding Association at Brunsberg, progeny testing, seed orchards, plus trees, tree banks and provenance tests. The third study tour had Malmö in southern Sweden as its starting point, the Chairman was Stig Wijkström, and Helge Johnsson the technical leader. The 64 participants were able to see the forest tree breeding station at Ekebo, provenance tests, progeny testing, and to visit the research station of the Swedish Match Company and the match factory at Jönköping.
The opening ceremony at the Folkets HUB in Stockholm took place on 23 August, and the Consultation then began regular meetings which went on until 30 August. The meetings were attended by 166 registered participants from 34 countries, and the international organizations represented, in addition to IUFRO, were the International Union of Biological Sciences (IUBS), the International Seed Testing Association (ISTA), the Biometrical Society, the Organization for Economic Co-operation and Development (OECD), and the East African Agriculture and Forestry Research Organization (EAAFRO).
King Gustav of Sweden graciously consented to be the High Patron of the Consultation. The Chairman of the Consultation and leader of the Swedish delegation was Erik Lundh, and the senior members of the Organizing Committee were:
Honorary member: Professor Nils Sylvén
Chairman: W. Plym Forshell
Vice-chairman: Professor Åke Gustafasson
General secretary: Helge Johnsson
Vice-secretary: C. Eklundh Ehrenberg
A panel of officers was submitted to and endorsed by the Consultation. For each section of the Consultation the officers concerned presided over the discussions in plenary meeting and conducted the work of the respective drafting committee. The combined findings of all the drafting committees were reported to the Consultation by the general rapporteur at the final plenary meeting.
A list of the consultation officers and other participants is given in Appendix E. Inauguration and opening speeches were given by W. Plim Forshell, Chairman of the Swedish Organizing Committee; Eric Holmquist, Swedish Minister of Agriculture; N. A. Osara, Director, Forestry and Forest Products Division, FAO; and J. D. Matthews, General Rapporteur of the Consultation and Senior IUFRO Representative.
The participants at the Consultation were asked by the Director of the Forestry and Forest Products Division of FAO to:
1. Review the present state and future prospects of forest genetics and tree improvement all over the world. This review is contained in the 11 chapters which form the major part of this final report.2. Propose a program for the closer co-ordination of development and wider spread of knowledge concerning forest genetics and tree improvement. This program is presented below as a series of recommendations, some addressed to governments, some to FAO and IUFRO.
3. Describe and endorse methods for mass-producing and distributing improved seed and plants for use in afforestation. This has been done in a series of technical recommendations.
In conclusion, there is no doubt that the Consultation was both successful and valuable. Our Swedish hosts worked hard and successfully to make the study tours instructive and memorable. There was a high standard of discussion during the section meetings in Stockholm and widespread agreement on most subjects. This final report is intended to provide a record of the Consultation: it is hoped that it will form a useful document for the promotion of forest genetics and tree improvement throughout the world.
GENERAL RECOMMENDATIONS
Recommendations to Governments
1. The regional timber trend studies published by FAO demonstrate the greatly increased demand for forest products in tropical countries, as a result of rising standards of living and rapid increases in population.
These trends, coupled with the generally low productivity of natural tropical forest stands, are focusing increased attention on the need to establish plantations of high-yielding species - either to supplement or replace the natural stands.
Despite a recent modest start in a few tropical countries, mainly on teak and tropical pines, the scale of forest genetics and tree improvement in the tropics is entirely inadequate, both in geographical distribution and species coverage, and bears no relation to its potential value and importance. All the indications are, however, that dramatic increases in production in many tropical species could be achieved by even the most elementary work on selection of seed trees, provenance research and formation of seed orchards. The major advantage in these regions is the speed with which results can be obtained.
The Consultation accordingly draws the attention o the governments of those tropical countries with forestry interests to the urgent need for developing planned programs of forest genetics and tree improvement. The Consultation also urges those organizations and countries concerned with multilateral or bilateral technical assistance in developing countries to consider thoroughly the possibility of including training centers and fellowships in forest genetics and tree improvement in their programs.
2. There is an urgent need for adequate information on forest genetics and tree improvement, and the Consultation proposes that these subjects should be included in the curriculum of schools, colleges and universities giving forestry courses. In many countries this may involve modification of the existing curricula, and the Consultation urges the governments and institutions concerned to take the necessary measures without delay.
3. The Consultation recognizes that choice of seed source is one of the main elements in the success and productivity of plantations, and that provenance studies provide a sound basis for the choice of seed sources. The Consultation recommends, therefore, that governments concerned with afforestation reforestation pay the greatest attention to the choice of seed sources and initiate or develop provenance research in close relation to their afforestation and reforestation programs.
4. The Consultation recognizes the importance to worldwide afforestation and reforestation programs of the acquisition of seeds from many sources of species growing in remote areas. The Consultation recommends, therefore, that forest geneticists and their governments work toward the establishment of seed collection offices. The Consultation commends the governments of Australia and Mexico for their help in collections of eucalypts and Mexican pines, and urges FAO to encourage and support the extension of such action to satisfy critical needs.
5. In numerous regions of the world natural or artificial stands representing valuable or proved sources of seed are in danger of disappearing or being hybridized with other races or species. The Consultation recommends, therefore, that the governments concerned take steps to assure the protection and conservation of these stands.
6. Just as the introduction of agricultural crop plants has been basic to the development of agriculture throughout the world, the introduction and testing of forest tree species is basic to the future success of forestry, especially when forestry is still in the pioneering stage as is often the case in countries at an early stage of development. Governments and research organizations should, therefore, co-operate in providing experimental lots of seed of native forest tree species (including representative geographic races) that have actual or potential value for planting in other countries. All possible efforts should be made toward setting up a service to establish representative provenances of such native species, collect seed of these provenances and make this seed available either free or at cost to approved research organizations or other government agencies (see also Recommendation 4).
7. The Consultation recommends that Member Nations of FAO and OECD should take steps to promote fully effective schemes for the certification of tree seeds and plants moving within their borders, thus paving the way for the certification of tree seed and plants moving in international trade. It welcomes, therefore, the initiative of OECD in forming an expert group to study and promote certification of such seeds and plants, also in view of the possibility for all Member States of FAO which are not members of OECD to become parties to seed certification schemes which the latter organization may recommend.
Recommendations to FAO and IUFRO
8. Widespread species of forest trees are of concern to many nations and research on such species will be most efficient if co-ordinated between nations by inter national organizations, such as FAO and IUFRO. Therefore, the Consultation asks Section 22 of IUFRO.
(a) to prepare and distribute summaries of the results obtained from past international provenance tests;(b) in liaison with FAO to organize new tests to extend the work on species previously studied and initiate new tests of species of broad interests, such as the eucalypts, Mexican pines, western North American conifers, teak and other tropical hardwoods, and Asian tree species;
(c) to prepare instructions concerning the collection of seeds, the design of field experiments and evaluation of results to aid in the development of local studies of provenance.
9. The Consultation recommends that Section 22 of IUFRO should form a Working Group on the Exchange of Poplar Seed for Provenance Research Purposes to provide minimum standards for the collection and inter national exchange of seed from poplar stands. This working group should act in close liaison with FAO's International Poplar Commission.
10. The introduction and testing of exotic forest trees is a truly international activity because the introduction of any species from one country to another concerns at least these two countries and it is generally of interest to a number of other countries as well. The Consultation agrees that the introduction and testing of exotic forest trees is a field of work in which an international organization such as FAO can render very valuable aid to forestry throughout the world, and particularly to countries at an early stage of development. Accordingly, the Consultation recommends that the Forestry and Forest Products Division of FAO should explore the possibility of setting up an advisory and information unit on the introduction of forest trees with actual or potential use as exotics. The functions of such a unit, to which the IUFRO sections concerned should render all possible assistance, might comprise:
(a) the provision of advice and information, in response to requests, on planning introduction programs, the species to be included, sources of seed, testing methods, properties of species and related subjects;(b) the dissemination of information on such tree species;
(c) recommendations for research to be undertaken by individual countries to fill gaps in the available information on these tree species.
Such a unit would need assistance from countries throughout the world to discharge these functions effectively and the Consultation recommends, therefore, that an Advisory Panel on the Introduction and Testing of Exotic Forest Trees should be established by FAO, possibly as a joint undertaking or at least in close association with IUFRO.
11. To promote the improvement of knowledge on tree physiology, the Consultation recommends that:
(a) IUFRO should establish a special working group on tree physiology;(b) FAO should support the organization of specific work in this field, and should consider the possibility of organizing a symposium on tree physiology in association with IUFRO.
12. The Consultation recognizes the present and future need in forest tree breeding for a great diversity of forest tree germ plasm, especially of rare or selected germ plasm, both natural and artificial, including seedling, clonal or hybrid material, and requests FAO to promote the establishment and maintenance of several germ plasm banks under the aegis of a suitable organization such as FAO itself.
13. The Consultation recommends that FAO should explore with IUFRO and other international or national agencies, such as the Commonwealth Forestry Bureau, the possibility of appointing a trained geneticist to examine the literature on the resistance of forest trees to parasites, accumulate the pertinent and accurate data on the genetic background of parasite resistance and have the review published.
14. The Consultation recognizes that resistance breeding requires the co-ordination of workers in the fields of forest genetics, pathology, entomology, and physiology, and therefore recommends that IUFRO should establish a working group on parasite resistance and improvement.
15. There is a need to study the regional patterns of flowering and fruiting in forest trees and this makes desirable the standardization of methods of measurement. The Consultation therefore recommends that a working group should be formed within Section 22 of IUFRO to make proposals for standardizing some of the measuring techniques applied in studies on the flowering and seed crop of forest trees.
16. The Consultation considers that the meeting in Stockholm has provided a concrete and satisfying example of the benefits of good collaboration between FAO and IUFRO The Consultation therefore recommends that this close collaboration between FAO and IUFRO should be strengthened in future so that FAO may draw on sound scientific advice and the work of the Sections of IUFRO can respond closely and rapidly to the economic and social needs of the time. The Consultation further recommends that the Permanent Committee of IUFRO should take account of the results of the Stockholm meeting and consider further collaboration with FAO in calling new consultations on other scientific and technical aspects of forestry.
17, The Consultation recommends that a Second Consultation on Forest Genetics and Tree Improvement should be held before the end of the current decade, and gratefully acknowledges the unofficial invitation made by the delegation of the United States of America.
TECHNICAL RECOMMENDATIONS
1. Both natural and induced mutants can be used in physiological, anatomical, and morphological investigations. Because they may be phenotypically different, these mutants also have value as genetic markers in inheritance studies.
2. Further work should be encouraged on the cytology of trees, and on precise chromosome mapping. With current techniques, it is difficult to determine accurately secondary constrictions in karyotype analyses but, as better techniques become available, seemingly similar karyotypes in different genera may well show marked differences in their chromosomes.
3. The genetic structure of populations should be investigated as thoroughly and defined as precisely as possible to permit realistic and reliable estimates of the genetic parameters (particularly heritabilities and the genetic correlation between characters) used in developing breeding programs. Tree breeders should be particularly careful when working with populations of mixed origin because of the possible occurrence of unpredictable genetic effects in such populations.
4. Progeny tests are vital to any breeding program. To obtain maximum information and to keep costs within tolerable limits, breeders should choose the mating design carefully. They should determine the BEST designs for field tests of progenies so that these field tests will also give reliable estimates of the breeding values of parents and the genetic parameters of the population. More uniformity trials should be made in planted stands to obtain information about the efficiency of different types of field tests. The results of such investigations and statistical data on field experiments should be published and made available to all tree breeders.
5. Research workers engaged in breeding for parasite resistance to disease and insect attack should make use of controlled pollination and progeny tests to determine precisely how the resistance is inherited (that is, whether due to major genes, genes with slight effects, dominant or recessive genes, or one, several or numerous genes). Open pollination progeny tests are not sufficient and should successively be replaced by progeny tests derived from controlled pollinations.
6. Attention should be paid to the interactions between genotypes and environment and to competition. Appropriate methods should be developed for their study.
7. Studies to develop methods for early testing or early diagnosis should be continued and intensified. They may soon provide the basis for faster progress.
8. Provenance research is costly of long duration. Studies to determine geographic patterns of inherent variation should, therefore, be emphasized when work is initiated on a new species. Preferably these studies should be undertaken before or at least concurrently with the establishment of seed source trials to test numbers of provenances in many localities.
9. More studies are needed of variation in wood properties within and between trees. A real danger is present, however, since some workers have freely extrapolated within-tree variation to between-tree variation. For example, conditions favoring fast growth of an individual tree usually also result in shorter fibers. This cannot be interpreted to mean that inherently fast growing trees are also short fibered trees, and such a relationship has not been proved. Yet many of the ideas regarding tree-to-tree differences in fiber characters have erroneously been extrapolated from within-tree data.
10. More studies are required to relate the effects of wood properties on the final product, both for solid wood products (saw-timber, veneer and others) and for uses such as pulp and chemically treated wood. Without better knowledge breeding for wood properties will proceed at a slow pace. Guidance and directives need to be given to the breeder so he can do his job efficiently. Prediction of needs in the distant future is involved.
11. Care must be taken to use enough trees in wood genetic studies so that results are sound and reliable. Many early reported results are merely indicators of trends and have errors so large that very little reliance can be placed on the absolute values obtained. However, the relative intensity of inheritance can be most helpful to the tree breeder.
12. For some species and some products specific gravity is high enough and wood properties other than specific gravity should be concentrated upon. Morphological characters of major importance are fiber length and fiber width.
13. Spiral grain is a wood character urgently needing investigation. One major problem involves sampling techniques. In the absence of definitive data, trees with spiral grain should be selected against in a breeding program, especially where the end product is sawtimber or veneer stock.
14. "Free splitting" is a major defect of some broadleaved species, especially in eucalyptus. It is not known whether this has a genetic basis and research is needed to determine its cause and control.
15. There is a dearth of genetic studies regarding chemical characters of wood. Such studies are urgently needed and may prove to be the most valuable of all work on the genetics of wood properties.
16. More genetic studies on wood of tropical broadleaved species are required, especially those of high value. Studies of the genetics of the wood properties of broadleaved species in general are scattered and insufficient.
17. The economic advantages of greater wood uniformity are stressed. There is not always need to "change" wood, but greater uniformity per se may be a major goal with real economic value.
18. It is probable that the more fundamental studies on the role of growth substances in flowering can best be done by biochemists working on herbaceous plants but tree physiologists and tree breeders should use their clones to do more work on the effects of environmental conditions on flowering. There is also good evidence of the importance of size of tree in the onset of flowering but the nature of this size effect requires careful study.
19. One promising treatment in seed orchard management is the application of fertilizers. Pruning of the crowns of the trees should, however, be used with care. Close attention should be given to the occurrence of stock-scion incompatibilities and the control of insect pests. In the long term, choice of rootstock and the use of growth substances to control flowering may be promising.
20. When clonal seed orchards are formed, the flowering and fruiting performance of the various clones must be studied and particularly the changes in flowering behavior with age, their responses to annual variations in climate, added nutrients and pruning. The efficiency of the seed orchards in producing regular crops of well-filled viable seed will depend to a large extent on the knowledge gained about the parent clones.
21. The great importance of inbreeding and associated phenomena in relation to breeding programs in general and seed orchards in particular is recognized as well as the fact that more must be learned about the variation in self-compatibility of the different parent clones and the role and nature of inbreeding. Evidence exists in Sweden and Australia of a good relation between the performance of the selfed and outbred progenies of certain parent trees.
22. The problem of mass-controlled pollination requires attention. One promising technique is the use of the polythene greenhouse developed in Finland as a very large isolation bag.
23. The importance of adequate identification of seed and plants and the value of good plantation records are stressed.
|
ai |
general combining ability of the i-th clone or line, etc. | |
|
A |
breeding value of an individual | |
|
b |
regression coefficient | |
|
CR |
correlated response to selection | |
|
DF |
degrees of freedom | |
|
E |
environmental deviation | |
|
E (MS) |
expectation of mean squares | |
|
F |
coefficient of inbreeding | |
|
G |
genotypic value | |
|
h² |
heritability | |
|
I |
index for selection | |
|
i |
intensity of selection | |
|
P |
phenotypic value | |
|
R |
response to selection | |
|
r |
correlation coefficient | |
|
sij |
specific combining ability of clones i and j | |
|
sA |
standard deviation of breeding values | |
|
V |
a variance component due to the source given by the subscript, thus: | |
|
|
VA |
component of genetic variance due to additive gene effects |
|
|
VD |
component of genetic variance due to dominance effects |
|
|
VE |
environmental variance |
|
|
VG |
genotypic variance |
|
|
VP |
phenotypic variance |
|
x |
a character to be selected for | |
|
y |
yield of a family, progeny, etc. | |
