Tree Physiology 15, 527--530 1995 Heron Publishing----Victoria, Canada

Seed orchards in development

G. B. SWEET
School of Forestry, University of Canterbury, Private Bag 4800, Christchurch, New Zealand Received March 11, 1994

Summary This paper reviews the role of seed orchards as output systems for genetic improvement programs. In this paper, technological changes since the 1950s are examined, with emphasis on recent developments. The need to equate the type of seed orchard to the type of forestry practiced is recognized, as is the significance of the relationship between seed orchards and other output systems for genetically improved material. Keywords: control pollination, genetic improvement program, management, open pollination, tree breeding.

Introduction Seed orchards are output systems for genetically improved material. Although they are not the only means of output, they are the most widely used. Whether seed orchards remain the major output system for genetic improvement programs (Giertych 1987) will depend on the biological and economic effectiveness with which they are able to transfer genetic gain from tree improvement programs to the forest. In this paper, I trace the development of seed orchards by reviewing principles rather than practices, with particular reference to conifers (see Faulkner 1975, Sedgley and Griffin 1989).

male and female strobili produced, and the differential timing of anthesis, individual clones do not contribute equally to the seed produced; that is, panmixis seldom occurs. Large numbers of clones are traditionally used to minimize the impact of self-pollination and parental imbalance. Second, the almost inevitable presence of pollen from outside the orchard substantially reduces genetic gains. Third, when ramets of individual clones are dispersed through the orchard, it is impractical to treat them individually to improve seed yields. Finally, management costs of orchards with tall trees are very high, and the time lag in getting genetically improved material to the production forest is considerable. For many species, pruning has not proved particularly effective. As a result of these limitations, modifications were proposed to the design of seed orchards and alternatives to seed orchards are being explored.

Downloaded from treephys.oxfordjournals.org by guest on March 10, 2011

Alternatives to seed orchards An obvious alternative to packaging improved genes in seeds is to distribute them in the cells of vegetatively propagated material. The many techniques of vegetative propagation offer considerable scope, first to take genetically improved material directly from the progeny test to the production forest, and second to increase the output from a clonal or seedling seed orchard. In both cases, the outcome is a form of clonal forestry, although true clonal forestry must involve clonal testing (Libby 1991). Clonal propagation techniques, which have been improving in effectiveness both biologically and economically for many years, may influence the nature of future output systems. In particular, techniques such as somatic embryogenesis, with their very high multiplication capacity, may modify the present role of seed orchards. Molecular biology techniques also have implications for seed orchards. Gene transfer has been accomplished successfully in several forest tree species and may become a standard component of tree improvement programs. Although it is unclear how gene transfer technology will be incorporated into existing breeding programs, the role of seed orchards as output systems for genetic improvement programs is likely to continue provided that desirable genes can be successfully transferred into all, or many, of the individual clones in an orchard.

Seed orchards Genetic improvement of forest tree species is usually based on recurrent selection for general combining ability. That is, the frequency of desirable genes in the population is increased progressively through cycles of selection and crossing (Shelbourne et al. 1989). Because this process involves sexual reproduction for progeny testing, and because most forest tree species are established with seedlings, seed has been the traditional output from genetic improvement programs. Seed orchards should represent an advancement, both in seed quality and quantity, over seed stands selected in high quality existing forests, because they are designed solely for seed production and are readily replaceable with orchards of more advanced genetic material. As initially conceived (Larsen 1956), seed orchards were clonal, but seedling seed orchards have also been established (e.g., White 1987). Clonal seed orchards can have considerable limitations (Sweet and Krugman 1977, Carson et al. 1992, AFOCEL 1992). First, as a result of clonal variability in the quantity of

528

SWEET

Improvements to conventional seed orchards If seed orchards are to remain the preferred output system, they need to remain competitive in several areas. (1) They should maintain the capacity to provide seed that optimizes genetic gains from the breeding program. This is likely to include provision for special purpose breeds and regional breeds, for which only relatively small seed quantities are required (Shelbourne et al. 1989). (2) They should by economically effective. Relevant factors of economic effectiveness include the time required for new-generational breeding material to come into production, and the cost of establishing isolation zones around open-pollinated seed orchards. (3) They should produce high quality output. The traditional Syrach Larsen orchard does not meet criteria (1) and (2) and thus improvements are needed. Among the first improvements to seed orchards were attempts to increase seed yields by physical and chemical improvements to soils, and by techniques such as root-pruning and girdling of trees (see papers in Faulkner 1975). Later, supplemental mass pollination (SMP) was used to increase seed yields, improve parental balancing and reduce the effect of outside wild pollens (e.g., Franklin 1971). In some Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) orchards, SMP was used in combination with water sprays, which delay orchard anthesis until after the bulk of contaminating outside pollens have been shed (e.g., Fashler and Devitt 1980). However, difficulties in applying these and the subsequently developed gibberellin application technologies (e.g., Pharis et al. 1987) to tall trees led to the development of a range of pruning and seed collection treatments that are still widely used. The pruning treatments are more successful in species with a 1-year seed development habit than in species with a 2-year development habit. However, even in species whose seed develops in 1 year, pruning normally removes the most important conebearing part of the crown. Sweet and Krugman (1977) proposed resolving the limitations of the conventional Pinus orchard by control-pollinating strobili on 3-m high ramets hedged in clonal blocks. The hedged ramets allow seed production treatments, pollination and other management operations to be applied easily from the ground. Placing ramets in clonal blocks enables treatments to be applied individually on a clonal basis. Block size can be adjusted for clonal fecundity. But most importantly, pollination can be controlled so that specific crosses can be made, with the potential for increases in genetic gain and for growing a range of different breeds on the same site. Control-pollinated orchards Although most seed orchards have not changed substantially since 1977, the New Zealand radiata pine (Pinus radiata D. Don) seed orchards were changed from open-pollinated (OP) to controlled-pollinated (CP) orchards in 1984 (Carson et al. 1992). Other single species orchards developed with similar objectives are HAPSOs (hedged, artificially pollinated seed orchards) and monoclonal orchards, both in Australia, and micro-orchards (trees planted in clonal rows and hand pollinated) in British Columbia and Georgia (Carson et al. 1992). The use of indoor breeding-halls (e.g., Adams et al. 1992)

offers similar potential. In terms of genetic quality, the CP orchards are superior to OP orchards, although SMP applied to individual whorls of flowers is a close substitute. However, CP orchards have not been widely adopted, possibly because of problems associated with increased conelet abortion and lower seed yields. Equally, however, Giertych (1987) has pointed out that, in many European countries, the major role of seed orchards is simply to make seed more readily available. Thus, there may be little incentive to establish CP orchards in many parts of the world. Orchard siting Recently, there has been considerable emphasis on the siting of seed orchards. There has been interest in transferring orchards in harsh climates toward warmer regions, to enhance quantity and earliness of flowering, seed maturation and isolation against foreign pollen (e.g., Enescu 1987). Long distance transfers of 1000 to 2000 km have been advocated for the latter reason (Koski 1987). However, there is increasing evidence for the existence of physiological aftereffects (Schmidtling 1987). These are effects of the climate at the orchard site on the climatic adaptability of the seedlings. The siting of seed orchards has also been used to optimize flower initiation. Sweet (1975) found that a more than 10-fold variation exists in cone production across radiata pine forest sites. For seed orchard management, high cone production and low vegetative growth is desirable, but little is known about the climatic and soil conditions under which this occurs. The use of potted grafts in a controlled greenhouse environment has been successful for small seeded hardwood species and some conifers (e.g., Sedgley and Griffin 1989). However, the significance of physiological aftereffects has not been widely assessed. With CP orchards, there is increasing interest in growing male and female orchards on separate sites. Seasonal earliness of anthesis then becomes important, because it may allow fresh rather than stored pollen to be used in a pollination program. In addition, ontogenetic earliness of flowering is key to the economic effectiveness of all orchards, because there appears to be a close correlation between heaviness of flowering and ontogenetic earliness. Physical management of orchards The traditional seed orchard is relatively straight forward to manage, apart from issues such as graft incompatibility (e.g., Copes 1970) and protection against fungal pathogens and insect pests (Bramlett 1987); however, the management of hedged orchards in clonal blocks is more complex. Much of the basic data needed to make effective management decisions does not exist but, as pointed out by Giertych (1987), the horticultural industry has considerable experience in intensive orchard management, much of which is relevant to forest tree seed orchards. In New Zealands CP orchards, it has been determined (e.g., Arnold 1990) that it is profitable to increase the stocking of ramets per hectare to the level of a meadow orchard (up to 10,000 ramets per ha). However, optimal meadow orchard

Downloaded from treephys.oxfordjournals.org by guest on March 10, 2011

SEED ORCHARDS IN DEVELOPMENT

529

stocking, in terms of seed production per hectare or returns per kg of seed, is not known, and returns from fertilizer, shelter or irrigation have not been evaluated. A pruning schedule has been developed for meadow orchards (Sweet 1992). There are particular difficulties in optimizing yield and financial return in orchards of Pinus, because pruning in the early years of a meadow orchard involves a constant trade-off between retaining existing immature cones and sacrificing them with the prospect of getting more cones in their place the next year. Quantitatively optimizing rotation age is also difficult.

Conclusions This article has concentrated on clonal seed orchards, but it should not be overlooked that determining the nature of output systems is an integral part of planning a breeding program (White 1987). The decision whether to use clonal or seedling seed orchards, or polycross seed orchards (Baradat et al. 1992) must be made on the basis of economic and genetic considerations. The principles established for clonal seed orchards are readily transferable to other types of orchard.
References Adams, G.W., J.C. Irving and M.S. Greenwood. 1992. Optimizations of environmental regimes for flowering in an indoor breeding hall for black spruce, white spruce and Jack pine. In Proc. AFOCEL/IUFRO Conference, Bordeaux, France. 1:219--227. AFOCEL. 1992. Mass production technology for genetically improved fast growing tree species. Proc. AFOCEL/IUFRO Conference, Bordeaux, France. Vol. 1, 441 p, Vol. 2, 532 p. Arnold, R.J. 1990. Control pollinated radiata pine seed----a comparison of seedling and cutting options for large-scale deployment. N.Z. For., November, pp 12--17. Baradat, P., C.E. Durel and P. Pastuszka. 1992. The polycross seed orchard: an original concept. In Proc. AFOCEL/IUFRO Conference, Bordeaux, France. 1:181--188. Bonnet-Masimbert, M. and P. Doumas 1992. Physiological approach to flowering induction in conifers. In Proc. AFOCEL/IUFRO Conference, Bordeaux, France. 1:51--59. Bramlett, D.L. 1987. Protection of pine orchards in the South-eastern United States. For. Ecol. Manage. 19:199--208. Carson, M.J., T.G. Vincent and A. Firth. 1992. Control-pollinated and meadow seed orchards of radiata pine. In Proc. AFOCEL/IUFRO Conference, Bordeaux, France. 2:13--20. Copes, D.L. 1970. Graft incompatibility and union formation on Douglas-fir. Silvae Genet. 19:101--106. Enescu, V. 1987. Climate and the choice of seed orchard sites. For. Ecol. Manage. 19:257--265. Fashler, A.M.K. and W.J.B. Devitt. 1980. A practical solution to Douglas-fir seed orchard pollen contamination. For. Chron. 56:237--241. Franklin, E.C. 1971. Pollen management in Southern seed orchards. In Proc. 11th South. Conf. For. Tree Improv., Atlanta, GA, USA, pp 218--223. Faulkner, R. 1975. Seed orchards. Forestry Commission Bulletin 54, 149 p. Giertych, M. 1987. Seed orchards in crisis. For. Ecol. Manage. 19:1--7. Koski, V. 1987. Long geographic transfers as a possible way of eliminating pollen contamination in advanced generation orchards of Pinus sylvestris. For. Ecol. Manage. 19:267--271. Larsen, C.S. 1956. Genetics in silviculture. Oliver & Boyd, Edinburgh, U.K., 224 p. Libby, W.J. 1991. Opening comments. In Proc. Clonal For. Workshop, Rotorua, New Zealand. FRI Bulletin No. 160, pp 11--12. Pharis, R.P., J.E. Webber and S.D. Ross. 1987. The promotion of flowering in forest trees by gibberellin A4/7 and cultural treatments: a review of the possible mechanisms. For. Ecol. Manage. 19:65--84. Rimbawanto, A., P. Coolbear and A. Firth. 1988. Artificial ripening of prematurely harvested cones of New Zealand Pinus radiata and its effect on seed quality. N.Z. J. For. Sci. 18:149--160. Schmidtling, R.C. 1987. Locating pine seed orchards in warmer climates: benefits and risks. For. Ecol. Manage. 19:273--283.

Chemical management of orchards Gibberellins A3 and A4/7 have been effective in enhancing the flowering of several genera and species in the Cupressaceae, Taxodiaceae and Pinaceae (Pharis et al. 1987). Paclobutrazol has also been identified as a reliable tool for increasing flower bud production in a range of species (AFOCEL 1992). In some orchards, chemical pollen emasculation is important. Because the optimal application time for chemicals is dependent on ramet size, clone, season and orchard location, it is desirable to develop a simple biological model that will integrate these variables (Bonnet-Masimbert and Doumas 1992).

Downloaded from treephys.oxfordjournals.org by guest on March 10, 2011

Pollination and pollen management in CP orchards With CP orchards, the opportunity exists to reduce clonal numbers and thereby increase genetic gains. In New Zealand, single parent pollination was initially associated with conelet abortion and low seed yields per cone. These difficulties have been overcome by the development of new methods for pollen extraction, storage and application technology (Sweet et al. 1992, Siregar 1994). Liquid pollination has proved effective in producing seeds that are larger and have higher germination capabilities than seed produced from normal (dry) pollinations (Setiawati 1994). There is considerable interest in carrying out liquid pollination without isolation, because it appears to be able to produce seed of the same genetic quality as obtained with isolation, but with considerable economic and logistic savings (Sweet et al. 1992). Also, it obviates the large reduction in seed yields per cone associated with the use of isolation bags (Setiawati 1994).

Physical seed quality There has been little emphasis in seed orchards on producing seed of a specific physical quality. Studies in New Zealand have shown that seed collected from 3-year-old meadow orchard ramets is comparable in size and germination capacity to seed collected from 6-year-old hedged ramets. The size of orchard seed can be increased by carefully timed irrigation treatments (Setiawati 1994). Rimbawanto et al. (1988) showed that by curing cones of P. radiata in a warm, low humidity environment, they could be harvested about 6 months before maturity and matured at a faster rate, which enables seed to be sown 12 months earlier.

530

SWEET Sweet, G.B. 1992. Seed orchard research and management in the 1990s----a New Zealand case study. In Proc. AFOCEL/IUFRO Conference, Bordeaux, France. 2:65--72. Sweet, G.B., R.L., Dickson, B.D. Donaldson and H. Litchwark. 1992. Controlled pollination without isolation----a new approach to the management of radiata pine seed orchards. Silvae Genet. 41:95--99. Sweet, G.B. and S.L. Krugman 1977. Flowering and seed production problems----and a new concept of seed orchards. In FAO/IUFRO 3rd World Consultation on Forest Tree Breeding, Canberra, Australia. 2:749--759. White, T.L. 1987. A conceptual framework for tree improvement programs. New For. 4:325--342.

Sedgley, M. and A.R. Griffin. 1989. Sexual reproduction of tree crops. Academic Press, London, U.K., 378 p. Setiawati, Y.G.B. 1994. Study of seed yield and physical quality in Pinus radiata D. Don seed orchards. M.S. Thesis. University of Canterbury, New Zealand, 152 p. Shelbourne, C.J.A., M.J. Carson and M.D. Wilcox. 1989. New techniques in the genetic improvement of radiata pine. Commonw. For. Rev. 68:191--201. Siregar, I.Z. 1994. Management of pollen, pollination and gibberellin A4/7 in Pinus radiata D. Don seed orchards. M.S. Thesis. University of Canterbury, New Zealand, 144 p. Sweet, G.B. 1975. Flowering and seed production. In Seed Orchards. Ed. R. Faulkner. Forestry Commission Bulletin 54, pp 72--82.

Downloaded from treephys.oxfordjournals.org by guest on March 10, 2011