1. 2000. Annual Report 1999/2000 - Pacific Northwest Tree Improvement Research Cooperative. Oregon State University, Oregon, USA. 31 p.

Keywords:      genetic tree improvement

                        tree/stand protection

                        tree physiology

                        reproduction

Abstract: Includes highlights of 1998-1999; a note to the cooperative members from Tom Adams; Introduction; Current research on seedling drought physiology of Douglas fir [Pseudotsuga menziesii], field drought study - genetics of drought sensitivity in older trees, early testing revisited, miniaturized orchard study, pollen contamination study; activities planned for 2000-2001; list of staff publications and abstracts; and a summary of financial support for the fiscal year 1999-2000.

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2.
Barclay, H.J. and Y.A. El-Kassaby. 1988. Selection for cone production in Douglas-fir adversely affects growth. In Proceedings: 10th North American Forest Biology Workshop: 'Physiology and genetics of reforestation', University of British Columbia, Vancouver, British Columbia, July 10-22, 1988. Eds. J. Worrall, J. Loo-Dinkins and D.P. Lester. pp. 149-151.

Keywords:      genetic tree improvement

                        genetic relationships

                        growth

                        reproduction

Abstract: Cone production and radial growth increment were studied for 8 years in a Douglas fir (Pseudotsuga menziesii) seed orchard on Vancouver Island, British Columbia, in 365 trees representing 29 open-pollinated (half-sib) families. Genetic correlations for the two traits were compared for each of the 8 years. Six of the correlations were significantly negative, while the remaining two were significantly positive. The predominantly negative correlation between cone production and growth indicates that selection for one of these two characters will effectively select against the other. The practice of selecting for high cone-producing trees in seed orchards may also be expected to yield slower-growing trees.

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3.
Colangeli, A.M., L. McAuley and J.N. Owens. 1990. Seasonal occurrence of potential ice-nucleating bacteria on Douglas fir foliage and seed cones. New-Forests 4(1): 55-61.

Keywords:      seed orchard management

                        tree/stand protection

                        tree/stand health

                        reproduction

Abstract: Plant frost damage can involve interactions between certain surface bacteria and low temperatures. The bacteria contain glycoproteins, which can nucleate ice above -5 degrees C, thus making the plants on which they live more susceptible to freezing. Preliminary studies to determine if bacteria were present on Douglas fir (Pseudotsuga menziesii), and whether they exhibited ice-nucleating properties, are reported. Total bacteria and fluorescent Pseudomonas populations were monitored on buds, conelets and foliage of five trees in a Douglas fir seed orchard on Vancouver Island, Canada, in April 1986 and between October 1986 and May 1987, over periods that spanned two pollination seasons. Seasonal variation in bacterial numbers was observed, with highest numbers occurring in late winter and early spring. Bacterial populations active in ice nucleation were found. Bacterial numbers during pollination were higher in 1986 than in 1987. Conelet abortion at pollination was also higher in 1986 (55%) than in 1987 (11%). A relation may exist between bacterial populations and conelet abortion at pollination.

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4.
Colangeli, A.M., J.N. Owens and S.J. Morris. 1989. Factors affecting cone and seed production in Douglas fir. BC Ministry of Forests FRDA-Report 057. 19 p.

Keywords:      seed orchard management

                        tree/stand protection

                        reproduction

Abstract: Reduced seed yield in 1986 on 4 Pseudotsuga menziesii trees in a British Columbia seed orchard was associated with inadequate pollination, low pollen vigour or viability, embryo abortion, and early ovule abortion. A study of bacterial populations suggested that there may be a causal relationship between this factor and conelet abortion.

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5.
Copes, D.L. and R.A. Sniezko. 1991. The influence of floral bud phenology on the potential mating system of a wind pollinated Douglas-fir orchard. Canadian-Journal-of-Forest-Research 21(6): 813-820.

Keywords:      genetic tree improvement

                        tree phenology

                        reproduction

                        genetic relationships

Abstract: Reproductive bud phenology was recorded from 1983 to 1989 in a Douglas fir (Pseudotsuga menziesii) clonal orchard near Monmouth, Oregon. Potential outcross efficiency was calculated for 20 clones from dates of male and female bud opening and pollination mechanism information. Potential outcross efficiency was limited to a maximum of 58% (1983) to 87% (1987). Cool weather before bud opening of the earliest clones delayed and compressed the breeding period and resulted in a greater percentage of trees having synchronous periods of pollen release and receptive seed strobili. Length of breeding season among years averaged 20 days and ranged from 16 to 27 days. Differences in phenology significantly affected the breeding system because the overall breeding period of the orchard clones exceeded the 8-day receptive period of individual clones by two or three times, and often prohibited or limited potential outcrossing between the earliest and latest clones. Outcrossing was greatest in clones with intermediate phenology and least in the earliest clones. The breeding system appears to be an almost continuous series of overlapping breeding subpopulations. Each year's breeding subpopulations were different from those of other years because of (1) large shifts in rank order of bud opening by 10 to 20% of the clones and (2) differences in the length of breeding season. Average temperature during March was linearly associated with time of floral bud opening. Geneticists may be able to use average temperature of the 4-week periods prior to opening of the earliest floral buds as a tool to identify seed crops formed during years with compressed breeding seasons. Such seed crops are potentially more diverse than seed crops produced during years with extended breeding seasons.

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6.
Copes, D.L. 1992. Effects of long-term pruning, meristem origin, and branch order on the rooting of Douglas-fir stem cuttings. Canadian-Journal-of-Forest-Research 22(12): 1888-1894.

Keywords:      seed orchard management

                        reproduction

Abstract: The rooting percentages of cuttings taken from 14 Douglas-fir (Pseudotsuga menziesii) clones were examined annually from 1974 to 1988. The trees were 10 and 13 yr old in 1974 and were pruned to 2.0 m in 1978 and 1979 and then recut annually to 0.5, 1.0, or 1.5 m, starting in 1983. The pruned trees showed no evidence of reduced rooting percentage even after 15 yr; average rooting increased from 47% in 1974 to 74% in 1986. Rooting percentage was significantly influenced by tree height. Cuttings collected from 0.5 m tall ramets exhibited better rooting than cuttings from 1.0 or 2.0 m tall ramets, and cuttings from 1.0 m tall ramets rooted better than cuttings from 2.0 m tall ramets. Rooting of cuttings collected from 0.5 m high subinterval zones within trees showed a negative linear relation between rooting percentage and collection height. Cuttings collected from the 0-0.5 m zone rooted 25% better than cuttings from the 1.5-2.0 m zone of the 2-m tall trees. A test of rooting of larger, more orthotropic cuttings gathered from the upper flat surface of pruned ramets indicated that the cuttings from the top rooted significantly less than the smaller, more plagiotropic cuttings from the contiguous side areas (24 vs. 33%, respectively). Meristems of secondary origin showed significantly greater rooting than meristems of primary origin. Comparison of rooting of second-order and first-order meristems of secondary origin indicated that second-order twigs averaged 26% better rooting than the first-order branch tips when the cuttings were collected in January and placed in the rooting beds in February.

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7.
Copes, D.L. and M. Bordelon. 1994. Effects of tree spacing and height reduction on cone production in two Douglas-fir seed orchards. Western-Journal-of-Applied-Forestry 9(1): 5-7.

Keywords:      seed orchard management

                        planting operations

                        reproduction

Abstract: Two treatments involving tree spacings (12 and 24 ft) within rows spaced 24 ft apart and height control (topped or not topped at 20 ft) were evaluated in 17- and 22-yr-old coast Douglas fir (Pseudotsuga menziesii var. menziesii) seed orchards in Oregon. Treatment differences for cone production were not significant because of the large tree-to-tree variation in cone production and, at one orchard, the confounding of location with treatment. Cone production in 1990 averaged slightly more than 2 bushels per tree (range 0 to 11.7 bushels). Average production per 96 linear ft of orchard row was 14.8 bushels.

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8.
Crook, R.W. and W.E. Friedman. 1992. Effects of pollen tube number and archegonium number on reproduction in Douglas-fir: significance for seed orchard management. Canadian-Journal-of-Forest-Research 22(10): 1483-1488.

Keywords:      genetic tree improvement

                        seed orchard management

                        reproduction

                        tree/stand health

Abstract: Analyses of the relations between pollen tube number or archegonium number and the number of fertilization events per ovule (fertilization number) in Douglas fir (Pseudotsuga menziesii) growing at a seed orchard near Centralia, Washington, indicated strong trends toward higher levels of simple polyembryony as both pollen tube number and archegonial number increased on a per seed basis. These relations have a significant bearing on the management of conifer seed orchards. Simple polyembryony has been proposed to be an effective means of increasing competition on a per seed basis in conifers and, potentially, the overall fitness of progeny. In conifers, supplemental mass pollination has the direct effect of increasing the number of pollen tubes per ovule. Clonal variation in average number of archegonia per ovule is also likely to exist among conifers. It is proposed that when used together, supplemental mass pollination and selection of clones with high archegonial averages may enhance the fitness of seed orchard progeny.

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9.
Edwards, D.G.W. and Y.A. El-Kassaby. 1988. Effect of flowering phenology, date of cone collection, cone-storage treatment and seed pretreatment on yield and germination of seeds from a Douglas-fir seed orchard. Forest-Ecology-and-Management 25(1): 17-29.

Keywords:      seed orchard management

                        reproduction

                        tree phenology

Abstract: The effects were studied of reproductive phenology, date of cone harvest, cone storage and seed pretreatment on yield and germination of seeds from a Douglas fir seed orchard in Victoria, BC, Canada. Flowering phenology (early, intermediate or late) had no discernible effect on seed maturation. Higher germination and yields of filled seeds were obtained from cones collected in mid-Aug., approximately 2 wk prior to cone opening, than from cones collected just as they began to open. Seeds extracted immediately following harvest germinated better than those from cones stored for 2 months. Seeds from all treatments were dormant and responded to prechilling by exhibiting increased germination rates. The implications of these findings for cone-crop management are discussed.

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10.
Edwards, D.G.W. and Y.A. El-Kassaby. 1995. Douglas-fir genotypic response to seed stratification. Seed-Science-and-Technology 23(3): 771-778.

Keywords:      seed orchard management

                        reproduction

                        genetic relationships

Abstract: Douglas fir (Pseudotsuga menziesii) responses to stratification duration were studied using wind-pollinated seeds from 15 seed-orchard clones, collected from a low elevation Douglas fir seed orchard in Saanichton, British Columbia, Canada. Germinative parameters (germination capacity, peak value, germination value, and germination rate and speed) were evaluated in response to four stratification periods (0, 3, 5, and 7 weeks). Significant differences among germinative parameters were observed indicating that the five-week stratification period represents the most appropriate treatment in minimizing variation caused by genetic differences. The results indicate that the International Seed Testing Association (ISTA) rules, which focus only on germination capacity, do not provide an adequate expression of seedlot dormancy, and since the rules are aimed at bulked seedlots, genetic differences, which can be large in heterogeneous forest tree seeds, are hidden. The results also demonstrate that extended stratification not only reduces the time in which seedlings become established, but also reduces seedling-emergence variation among parental lines.

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11.
El Kassaby, Y.A. 1995. Evaluation of the tree-improvement delivery system: factors affecting genetic potential. Tree Physiology 15:545-550.

Keywords:      genetic tree improvement

                        nursery operations

                        seed orchard management

                        genetic relationships

                        reproduction

Abstract: Possible causes of the genetic erosion that occurs during the fragmented phases of the tree-improvement delivery system (a term used for the domestication process in forest trees) are reviewed. The impacts of intentional and unintentional directional selection during phenotypic selection, seed production (with its associated reproductive-phenology asynchrony, fecundity differential and varying propensity to inbreeding), seed processing and storage, and seedling production are evaluated. Allozyme analysis data were used to compare heterozygosity of first-generation seed orchards of western red cedar (Thuja plicata), Sitka spruce (Picea sitchensis) and Douglas fir (Pseudotsuga menziesii) with that of their corresponding natural populations. In general, genetic diversity and heterozygosity parameters of seed orchards are higher or similar to those observed in their natural-population counterparts. However, parental contribution to the resultant seed orchard seed crops is consistently asymmetrical, and this is a major cause of genetic erosion. In most cases, less than 20% of an orchard's clones contribute 80% of the cone crop, thus reducing the effective population size. Because seed germination of coniferous tree species is under strong maternal genetic control, the combined effects of differences in reproductive output and germination, as well as of management practices (e.g., simulated long-term storage of seed showed that loss of viability during storage is genotype specific), cause unintentional directional selection during seedling production. This review confirms the need for genetic monitoring of each phase of the tree-improvement delivery system, so that practical solutions can be developed to alleviate genetic erosion.

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12. El Kassaby, Y.A. 2000.
Representation of Douglas-fir and western hemlock families in seedling crops as affected by seed biology and nursery crop management practices. Forest Genetics 7(4): 305-315.

Keywords:      genetic tree improvement

                        nursery operations

                        genetic relationships

                        tree/stand health

                        reproduction

Abstract: The impact of container-nursery management practices on the genetic representation of seedling crops of Douglas fir (Pseudotsuga menziesii) and western hemlock (Tsuga heterophylla) were evaluated. Two experiments, one for each species, were conducted in British Columbia, Canada, to determine the cumulative effects of seed-donor variation on germination parameters (percent and speed) and their interaction with container-nursery practices of thinning and culling on the genetic representation of each seed-donor in the resultant seedling crops. The experimental work was conducted on seedlots that were represented with equal contribution of seeds from 15 seed orchard parents (families). In each experiment, a total of 25 920 seeds were sown in four different arrangements to compare the crop development under single-, two-, and three-seeds per cavity (seeds within cavity were selected randomly among the 15 families) and family blocks (seeds within block belonged to one family). This experimental design allowed determination of inter- and intra-family competition. Within each experiment, a total of 15 015 cavities were used and the identity of every seed within every cavity within each arrangement was maintained throughout the study. Families were compared based on: (1) changes in their rank order from seedling emergence (germination) to post-thinning and post-culling status, and (2) relative performance of each family from seed contribution to seedling production. Changes were observed in both assessments (i.e., rank and relative contribution). Path analysis was used to determine the percent contribution of each factor to seedling production. It was determined that seedling germination, germinant thinning, and seedling culling all affected seedling production, indicating the presence of several consecutive unintentional bottlenecks in the process. Family sowing with culling standards that recognize the growth differences among families in the nursery and single seed sowing after understanding the inter-/intra-family competition are recommended for seedling production to maintain seedling-crop family representation.

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13. El Kassaby, Y.A., S. Barnes, C. Cook and D.A. MacLeod.
1993. Supplemental mass pollination success rate in a mature Douglas-fir seed orchard. Canadian-Journal-of-Forest-Research 23(6): 1096-1099.

Keywords:      genetic tree improvement

                        seed orchard management

                        reproduction

Abstract: Supplemental mass pollination (SMP) success rate in a mature Douglas fir (Pseudotsuga menziesii) seed orchard in British Columbia was studied with the aid of a unique electrophoretically detectable allozyme marker. Four SMP methods were tested during periods of maximum pollen release in 1990: operational applications of pollen (pollen applied once to a branch at average receptivity), and branch applications that were carried out during one, two, or three visits (a first pollen application at maximum receptivity and subsequent applications 2 and 4 days later) were conducted. No significant increase in seed-yield traits (total number of seeds per cone, number of filled seeds per cone and seed efficiency) was observed between wind-pollinated and SMP-treated cones. However, multiple branch visits showed significant increases in SMP success rate (18%) when compared with the operational visit (8%) or the one branch visit (9%) results. No significant increase in SMP success rate was obtained when the number of branch visits was increased from two (17.8%) to three (17.9%). It was concluded that operational SMP has the potential to improve the genetic value of the treated Douglas fir crops even under high pollen load. The degree of improvement is dependent directly on SMP fertilization success and indirectly on the genetic value differential between the supplemented and the orchard's ambient pollen, the frequency of application, and the quality (i.e. viability) of the pollen applied.

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14. El Kassaby, Y.A., K. Chaisurisri, D.G.W. Edwards and D.W. Taylor.
1993. Genetic control of germination parameters of Douglas-fir, Sitka spruce, western redcedar, and yellow-cedar and its impact on container nursery production. In Dormancy and barriers to germination. Proceedings of an international symposium of IUFRO Project Group P2.04-00 (Seed problems), Pacific Forestry Centre, Victoria, British Columbia, Canada. Ed. D.G.W. Edwards. pp. 37-42.

Keywords:      genetic tree improvement

                        nursery operations

                        genetic relationships

                        reproduction

Abstract: The genetic control of germination parameters (germination capacity, peak value, and germination value) in Douglas-fir (Psuedotsuga menziesii (Mirb.) Franco), Sitka spruce (Picea sitchensis (Bong.) Carr.), western redcedar (Thuja plicata Donn), and yellow-cedar (Chamaecyparis nootkatensis (D.Don) Spach.) was studied using wind-pollinated seeds collected from several seed orchards. The extent of genetic control over rhese parameters was assessed through the determination of broad-sense heritabilities. The impact of genetic control of these parameters on the expected genetic diversity of container nursery seedling crops is evaluated.

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15. El Kassaby, Y.A. and C. Cook.
1994. Female reproductive energy and reproductive success in a Douglas-fir seed orchard and its impact on genetic diversity. Silvae-Genetica 43(4): 243-246.

Keywords:             genetic tree improvement
                               genetic relationships
                               reproduction

Abstract: The relationship between reproductive energy and reproductive success and its impact on clonal female gamete contribution were studied in a clonal Pseudotsuga menziesii seed orchard. The effect of among- vs. within-clonal variation on seed-cone crop (energy) and filled-seed yield (success) was determined by partitioning their respective variance components. Clonal gametic representations were expressed using Griffin's [Australian Forest Research (1982) 12, 295-302] parental-balance curves and Crow and Kimura's [An introduction to population genetic theory (1970) New York, USA; Harper and Row Publishers] effective population number. Seed-cone and filled-seed yields produced similar parental-balance curves and female effective population numbers. Although similar parental-balance curves and female effective population numbers were produced, the actual clonal gametic representation differed when the contribution of any set of specific clones were considered. It was concluded that parental-balance curves and female effective population numbers provide a static description of the genetic representation and do not illustrate the dynamics of clonal reproductive outputs.

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16. El Kassaby, Y.A., D.G.W. Edwards and C. Cook.
1990a. Impact of crop management practices on seed yield in a Douglas-fir seed orchard. Silvae-Genetica 39(5-6): 226-230.

Keywords:      seed orchard management

                        tree/stand protection

                        reproduction

                        tree/stand health

Abstract: The impact of two crop-management practices, supplemental mass pollination (SMP) and overhead cooling, on seed yield in a 13-yr-old Douglas fir (Pseudotsuga menziesii) seed orchard was studied in Saanichton, British Columbia. A 2x2 factorial trial of SMP/no SMP and cooling/no cooling was applied. There were no significant differences in potential seed yield per cone, average number of successful fertilizations, and average number of filled seeds per cone between cooling or SMP treatments or combinations. Results indicated that within-orchard pollen cloud was not a factor limiting seed yield. Average number of seeds infested by the Douglas fir seed wasp (Megastigmus spermotrophus) larvae was significantly (P<0.05) less when cooling was applied, indicating that the treatment was effective in disrupting the synchrony between the presence of ovipositing females and developing cones.

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17. El Kassaby, Y.A., D.G.W. Edwards and D.W. Taylor.
1990b. Effect of water-spray cooling treatment in a Douglas-fir seed orchard on seed germination. New-Forests 4(2): 137-146.

Keywords:      seed orchard management

                        reproduction

                        tree phenology

Abstract: As part of an assessment of seed orchard crop management practices, results are given of an experiment undertaken to test the effect of cooling treatment on germination in a 13-year-old full-sib Douglas fir (Pseudotsuga menziesii) seedling seed orchard at Saanichton, British Columbia, Canada. Overhead cooling by sprinklers during February-March, used in order to delay flowering, produced non-significant effects on germination capacity, germination rate, germination value, and abnormal germination of seeds. The percentage of variation accounted for by cooling treatment ranged from 0.0 to 1.0%. In contrast, variation among trees within each treatment (i.e. cooled or not cooled) ranged from 28 to 46%. These effects were significant, indicating that each tree has an individual germination pattern. The temporal delay of reproductive phenology caused by temperature manipulation as a result of the cooling treatment was judged to be within the species' biological limits.

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18. El Kassaby, Y.A., A.M.K. Fashler and O. Sziklai.
1984. Reproductive phenology and its impact on genetically improved seed production in a Douglas-fir seed orchard. Silvae-Genetica 33(4/5): 120-125.

Keywords:      seed orchard management

                        tree phenology

                        reproduction

Abstract: Reproductive phenology of buds was monitored for 2 yr in a Douglas fir seed orchard to determine the validity of the theory of panmictic equilibrium. There was a significant variation within clones and families in dates of seed-cone and pollen-cone bud burst, suggesting a major deviation from panmictic equilibrium. This would reduce the size of the breeding population and reduce seed yield. This effect could be reduced, and seed production maximized, either by the use of an irrigation mist system to delay bud development, or by the intensive application of booster pollination.

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19. El Kassaby, Y.A., J. Parkinson and W.J.B. Devitt.
1986. The effect of crown segment on the mating system in a Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seed orchard. Silvae-Genetica 35(4): 149-155.

Keywords:      genetic tree improvement

                        seed orchard management

                        genetic relationships

                        reproduction

Abstract: The mating system in a mixed clonal/seedling Douglas fir seed orchard was studied by analysing 6 polymorphic allozyme loci. Trees were subjected to cooling treatment to retard bud development and compact the pollination period. The outcrossing rates of upper and lower crown levels and northern and southern crown aspects were estimated and compared for clonal and seedling material separately using single and multilocus methods. Single locus estimates of outcrossing rate ranged from 0.645 to 0.999 and were significantly heterogeneous. No significant differences between the multilocus estimates were observed between crown levels or aspects for either the clonal or seedling material. For all comparisons, the unweighted or weighted single locus means were lower than those obtained by the multilocus method, indicating the presence of other types of consanguineous mating in addition to selfing. In general, higher outcrossing rates were observed in the clonal trees than in those derived from seedlings. The implications of these results for seed orchard management are discussed.

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20. El Kassaby, Y.A. and K. Ritland.
1986. The relation of outcrossing and contamination to reproductive phenology and supplemental mass pollination in a Douglas-fir seed orchard. Silvae-Genetica 35(5/6): 240-244.

Keywords:      genetic tree improvement

                        seed orchard management

                        tree phenology

                        genetic relationships

                        economics

                        reproduction

Abstract: A study was made using allozyme markers of outcrossing and contamination rates in relation to reproductive phenology and supplemental mass pollination in a Douglas fir seed orchard in British Columbia, Canada. Supplemental mass pollination was applied only to the intermediate reproductive phenology class, which showed a high outcrossing rate and no contamination. Both early and late reproductive phenology classes showed significant contamination, but the outcrossing rate for the former was significantly higher than for the latter. These results show that interpretation of seed crop genetic quality based on outcrossing alone could be misleading. The rate and source of contamination, reproductive phenology and crop size should also be considered. The practicability and economics of supplemental mass pollination in avoiding both selfing and contamination are discussed. It was concluded that waterspray cooling and/or supplemental mass pollination of early and late reproductive phenology classes in moderate or good cone-crop years would be an effective management practice. The option of aborting small cone crops in mature orchards is also discussed.

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21. El Kassaby, Y.A. and A.J. Thomson.
1996. Parental rank changes associated with seed biology and nursery practices in Douglas-fir. Forest Science 42:228-235.

Keywords:      genetic tree improvement

                        nursery operations

                        reproduction

                        genetic relationships

                        tree/stand health

Abstract: The impact of container-nursery management practices on the genetic composition of seedling crops was evaluated in an experimental study in a seed orchard in British Columbia. The study was designed to determine the cumulative effects of: (1) differences in parental reproductive output in bulk seedlots; (2) parental variation in germination parameters (percentage and speed); and (3) the interaction of these parameters with container-nursery practices of thinning and culling, and their effect on the genetic representation of parents in the resultant seedling crops. Results from the experimental study were compared with predictions of a stochastic simulation designed to estimate the consequences of differential parental seed contribution, and seed germination percentage and germination speed on indicators of crop performance. The experimental study was conducted on a Douglas-fir (Pseudotsuga menziesii) bulk seedlot that was representative of the differential seed contributions from 19 seed orchard parents. The nursery study included a total of 42 000 seeds. Seeds were sown at three seeds per cavity. Within the 14 000 cavities used, the identity of every seed was maintained throughout the study. Comparisons of parents were made based on: (1) changes in their rank order from sowing to postthinning and postculling status; and (2) relative performance from seed contribution to seedling production. Changes were observed in both assessments. Path analysis was used to determine the percentage contribution of each factor to seedling production. It was determined that germination, thinning, and culling contributed 66, 20, and 14%, respectively, to seedling production, indicating the presence of three consecutive bottlenecks in the process. Single seed or individual family sowing in the nursery was recommended for seedling production to maintain genetic diversity.

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22.
Fashler, A.M.K. and Y.A. El-Kassaby. 1987. The effect of water spray cooling treatment on reproductive phenology in a Douglas-fir seed orchard. Silvae-Genetica 36(5-6): 245-249.

Keywords:      genetic tree improvement

                        seed orchard management

                        tree/stand protection

                        genetic relationships

                        reproduction

                        tree/stand health

Abstract: The effectiveness of reproductive bud cooling on genetic efficiency in a Douglas fir seed orchard in British Columbia, Canada, was tested by comparing the reproductive bud phenology in 3 cooled and 3 uncooled years. The cooling system was found to affect 2 major elements affecting seed orchard genetic efficiency, namely pollen contamination levels and panmictic equilibrium, as well as insect infestation, frost damage, seed yield and management effectiveness. Based on these results, a permanent irrigation/cooling system is recommended for Douglas fir seed orchards.

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23.
Fernando, D.D., J.N. Owens and P.v. Aderkas. 1998. In vitro fertilization from co-cultured pollen tubes and female gametophytes of Douglas fir (Pseudotsuga menziesii). Theoretical-and-Applied-Genetics 96(8): 1057-1063.

Keywords:      genetic tree improvement

                        reproduction

Abstract: A previous attempt on in vitro fertilization (IVF) in conifers resulted in pollen tube penetration of female gametophytes, but because of the rapid decline in egg viability, no further interaction occurred. It is reported that, for the first time, IVF has been achieved in conifers. Using Douglas fir (Pseudotsuga menziesii), a two-step process is described which involved induction of pollen tubes in culture followed by introduction of isolated female gametophytes at the tips of growing pollen tubes. Pollen tubes penetrated the introduced isolated female gametophytes at various places, but a number of tubes entered the egg cell through the neck cells similar to the in vivo condition. Under current culture conditions, longevity of pollen tubes and eggs was improved resulting in the release of sperms, fusion of gametes, and initial formation of the proembryo. Continued plasmolysis of the egg limited the number of successful gametic interactions. IVF has been accomplished in flowering plants in several ways, but the gametophyte-gametophyte IVF system described in here is unique. IVF offers a novel breeding technology that takes advantage of the sexual reproductive route. When coupled with hybridization and genetic transformation, IVF could result in the development of stable novel genotypes of economically superior trees.

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24.
Fernando, D.D., J.N. Owens, P.v. Aderkas and T. Takaso. 1997. In vitro pollen tube growth and penetration of female gametophyte in Douglas fir (Pseudotsuga menziesii). Sexual-Plant-Reproduction 10(4): 209-216.

Keywords:      genetic tree improvement

                        reproduction

Abstract: Pollen tube and female gametophyte interactions in Douglas fir (Pseudotsuga menziesii) were examined in vitro. Formation of pollen tubes in Douglas fir occurred on a modified MS medium in which concentrations of H3BO3 and Ca(NO3)2 were altered and supplemented with sucrose and polyethylene glycol. Addition of 100 micro g/ml H3BO3 and 300 micro g/ml Ca(NO3)2 resulted in optimum pollen viability. Lack of H3BO3 inhibited pollen tube formation. Addition of H3BO3 and Ca(NO3)2 significantly increased pollen tube formation within one week in culture. Using a medium supplemented with mannitol, viability of Douglas fir pollen can be sustained for 7 weeks in culture, about the same length of time as in vivo. However, pollen tubes are not formed. This suggests that the factors responsible for tube formation reside in the external environment of the pollen. Culture of female gametophytes to examine egg viability and longevity had not been done previously. Egg viability in culture is short-lived, and therefore the window to study and manipulate events of fertilization in Douglas fir is very limited. In spite of this, about 7% of the female gametophytes that were co-cultured became penetrated by pollen tubes. In vitro archegonial penetration has been repeatedly achieved, but pollen tubes also penetrated other parts of the female gametophytes. Pollen tubes also penetrated non-viable eggs. Most female gametophytes were not penetrated because of pollen tube branching and swelling, failure of tubes to orient towards the female gametophytes, or premature pollen tube death due to plasmolysis. This report outlines the first attempt towards in vitro fertilization in conifers.

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25.
Johnson, D.R., G.P. Markin, R.C. Reardon and W.K. Randall. 1984. Injecting Metasystox-R at three spacing intervals to improve seed yield in Douglas-fir. Journal-of-Economic-Entomology 77(5): 1320-1322.

Keywords:      tree/stand protection

                        reproduction

Abstract: Injecting oxydemeton-methyl into Douglas firs (Pseudotsuga menziesii) using the Mauget Inject-A-Cide technique at 3 spacing intervals in studies in western Oregon in 1982 significantly reduced populations of Contarinia oregonensis and increased filled seed yield when compared with untreated controls. Injectors were placed at intervals of 5, 10 or 15 cm of circumference at breast height and delivered 0.1 g a.i./cm of circumference. The results did not differ among the treatments.

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26.
Koerber, T.W. and G.P. Markin. 1984. Metasystox-RReg. injections increase seed yield of Douglas-fir in California, Oregon, and Washington. In Proceedings of the cone and seed insects working party conference, Working Party S20701, Southeastern Forest Experiment Station, Asheville, NC. Ed. H. Yates, III. pp. 137-146.

Keywords:      tree/stand protection

                        tree/stand health

                        reproduction

Abstract: Injections of 1.5 g for each 15 cm of tree girth reduced seed damage caused by Contarinia oregonensis and Barbara colfaxiana, but not that by Megastigmus spermatrophus [M. spermotrophus]. On sites with insect populations high enough to cause substantial seed losses, treatment increased seed yield per cone by 38-162%.

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27.
Krakowski, J. and Y.A. El-Kassaby. 2003. Effects of stratification and simulated aging on germination of Douglas-fir seed from a clonal seed orchard. Forest-Genetics 10(1): 65-70.

Keywords:      nursery operations

                        tree/stand health

                        reproduction

Abstract: Seeds from 15 Douglas-fir (Pseudotsuga menziesii) clones were germinated in a factorial design with two pre-treatments (unstratified and stratified) and seven simulated aging periods (0, 2, 4, 7, 10, 12 and 14 days). Simulated aging consisted of high temperature (40 degrees C) and relative humidity (100%) exposure, which simulates physiological stresses and consequent deterioration in long-term storage. Seed deteriorated as aging treatments lengthened; no germination occurred after 12 days. Germination parameters (capacity, peak value, speed, completeness) were calculated, and pre-treatment and aging effects evaluated using a mixed model analysis of variance. Germination completeness and speed were higher after two days of aging for stratified seed, whereas only peak value increased for unstratified seed. After four days aging, all parameters decreased. Two days of aging enhanced germination capacity of unstratified seed by 15%, but stratified seed was still 13% higher. Douglas-fir seed should be stratified before germination, but unstratified seed can be exposed to 40 degrees C and 100% humidity for two days to augment seedling stock during the growing season. Ex situ Douglas-fir genetic resource conservation, as well as more adequate representation of planted genotypes across the landscape, can benefit from two days of aging, which would ensure slowly-germinating genotypes are represented in the population.

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28
. Malavasi, M.d.-M., S.G. Stafford and D.P. Lavender. 1985. Stratifying, partially redrying and storing Douglas-fir seeds: effects on growth and physiology during germination. Annales-des-Sciences-Forestieres 42(4): 371-383.

Keywords:      nursery operations

                        reproduction

                        growth

Abstract: Douglas fir seeds collected from one coastal and one inland source in Oregon, USA, were stratified at 3 degrees C for 28 days at 45% m.c., and then redried to 35% or 25% m.c. and/or stored at 3 degrees C for 1 or 3 months. Redrying stratified seeds to 35% m.c. did not affect the m.c. of embryos or gametophytes, but redrying to 25% m.c. reduced the m.c. of all seed structures. Three months storage did not alter moisture distribution within seeds. Stratification reduced the germination % of seeds from the inland source, but hastened germination speed of seeds from both sources. Redrying stratified seeds to 35% and 25% m.c. increased seed vigour and seedling length and dry weight. Storing stratified seeds without redrying them generally reduced seed vigour. These results suggest that it would be advantageous to redry seeds to a range of 25-35% m.c. directly before sowing in order to produce vigorous seedlings or allow expression of stratification benefits.

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29.
Miller, G.E. 1983b. When is controlling cone and seed insects in Douglas-fir seed orchards justified? Forestry-Chronicle 59(6): 304-307.

Keywords:      seed orchard management

                        tree/stand protection

                        economics

                        reproduction

Abstract: Two seed orchards in British Columbia were sprayed with dimethoate in 1981, and the costs of estimating crop size and insect infestation and of dimethoate application were recorded. The cost/tree was $2.31 or $3.68 (including sprayer rental). Benefit/cost ratios were calculated and plotted against number of cones/tree and varying increases in yield (3-24 filled seeds/cone) due to protection. The number of cones a tree must bear and the increased yield/cone required to cover the cost of one dimethoate application are given in graphs for seed values of $150-1000/kg.

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30.
Miller, G.E. 1986. Damage prediction for Contarinia oregonensis Foote (Diptera: Cecidomyiidae) in Douglas-fir seed orchards. Canadian-Entomologist 118(12): 1297-1306.

Keywords:      seed orchard management

                        tree/stand protection

                        tree/stand health

                        reproduction

Abstract: Damage to Douglas fir (Pseudotsuga menziesii) in British Columbia at cone harvest by Contarinia oregonensis was positively correlated with the number of egg-infested scales per conelet in the spring. Reducing the average number of galled seeds per cone by 1.5 increased the average number of filled seeds per cone by 1.0 in insecticide trials. Optimum sample sizes for estimating average densities of egg-infested scales were calculated to be 1 conelet/tree and 150 trees/orchard. The mean crowding variable was linearly related to average density, so a sequential sampling technique relative to a critical density was developed for determining the need for control measures.

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31.
Minore, D. 1986b. Germination, survival and early growth of conifer seedlings in two habitat types. Pacific-Northwest-Forest-and-Range-Experiment-Station,-USDA-Forest-Service Research-Paper PNW-RP-348. ii + 25 p.

Keywords:      planting operations

                        reproduction

                        tree/stand health

                        growth

Abstract: Seed germination, and seedling survival and early growth of Douglas fir, western hemlock, Abies procera and A. amabilis were studied on clear felled sites in the A. amabilis/Achlys triphylla and A. amabilis/Vaccinium membranaceum/Xerophyllum tenax habitat types in the McKenzie River basin, Oregon. Severe seed predation and high surface soil temp. in both habitat types resulted in almost total seedling mortality during the first season when seeds had been sown on a few large, consolidated seedbeds. Seedling survival was n.s.d. between habitat types. Survival of Douglas fir (the only species tested) was better when seeds were sown in small scattered spots. Habitat type and addition of forest humus did not affect survival and growth, but shading by stumps and protection from predation using plastic berry baskets were both beneficial. Soils from both habitats had similar colour, texture, pH and nutrient content. In greenhouse studies, seeds of all species began germinating earlier and continued to germinate for longer in soil from Achlys sites than in soil from Xerophyllum sites. It is suggested that unmeasured biological activity may have been responsible for differences in germination activity between the two soil types.

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32.
Minore, D., H.G. Weatherly and P.G. Cunningham. 1993. Sowing at 1.5-cm (0.6-inch) depth produces heaviest Douglas-fir roots in small containers. Tree-Planters' Notes 44(3): 122-124.

Keywords:      nursery operations

                        tree morphology

                        growth

                        reproduction

Abstract: Sowing seeds of Douglas fir (Pseudotsuga menziesii) at five depths in Leach Super Cells indicated that the only benefit of deep sowing in small containers occurred at a depth of 1.5 cm. Sowing at this depth produced heavier roots without a significant reduction in seedling emergence. Sowing at greater depths significantly reduced seedling emergence and growth.

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33.
Mohammed, G.H., K.R. Patel and W.E. Vidaver. 1989. The control of adventitious root production in tissue-cultured Douglas-fir. Canadian-Journal-of-Forest-Research 19(10): 1322-1329.

Keywords:      nursery operations

                        reproduction

                        growth

                        tree morphology

Abstract: Rooting percentage and root number in tissue-cultured Douglas fir (Pseudotsuga menziesii) were examined to assess the influence of rooting substrate, the concentrations of sucrose and boron in the rooting medium, shoot height, and shoot generation. Peat/perlite was a better substrate than agar, producing 70% compared with 0% rooted shoots, respectively. On peat/perlite, cell divisions were organized and were associated with tracheid nests, whereas on agar proliferation was neither organized nor restricted to the nests. An optimum sucrose concentration of 4% was found for the production of nodular or rooted shoots. At 4% sucrose and 3 mg/litre boric acid, 100% of the shoots rooted, and the mean root number was 11. Rooting percentage and root number were significantly greater with shoots that were 3 cm tall rather than 2 or 1 cm tall. Shoot responses were more rapid in third and fourth generation shoots, with at least 80% rooted or nodular after 4 weeks, compared with only 36% from the second generation.

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34.
Muller, C., E. Falleri, E. Laroppe and M. Bonnet Masimbert. 1999. Drying and storage of prechilled Douglas fir, Pseudotsuga menziesii, seeds. Canadian Journal of Forest Research 29:172-177.

Keywords:      nursery operations

                        reproduction

Abstract: Douglas fir, Pseudotsuga menziesii, seeds exhibit relative dormancy as they do not germinate at suboptimum temperature (15°C), whereas at optimum temperature (20°C) some germination occurred. Thus, germination at 15°C was chosen to estimate dormancy release. In the first experiment, seeds were prechilled at 32% moisture content (MC) for 0-34 weeks at 3°C. Long chilling treatments enhanced germinability and, more markedly, germination speed both at 20°C and at 15°C. Seeds pretreated for the longest periods were then dried to 6.7% MC and stored up to 6 months without any detrimental effect on germination at 15°C. In the second experiment, seeds from a second seedlot were prechilled for 18 weeks and then stored at three different MCs (6.7, 7.2 and 8.1%) over a period of 17 months. Seeds stored at the lowest MC germinated fastest and to the highest percentage both at 15 and 20°C. In the nursery, seedling emergence tests confirmed results from this experiment.

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35.
Poulsen, K.M. 1996. Prolonged cold, moist pretreatment of conifer seeds at controlled moisture content. Seed Science and Technology 24:75-87.

Keywords:      nursery operations

                        reproduction

                        tree/stand health

Abstract: Cold pretreatment of imbibed seed (stratification) and seed kept at controlled moisture contents for 3-14 weeks was tested for commercial seedlots of Abies nordmanniana, Abies procera, Pseudotsuga menziesii and Picea sitchensis in order to give recommendations for nursery practice. The temperature tolerance of treated seed was tested by germinating seed at four temperature regimes. It was possible to pretreat Abies nordmanniana seed at controlled moisture content, but the performance was not improved compared to the traditional six weeks stratification of imbibed seed. For Abies procera seed pretreatment at controlled moisture content for 2 + 8 weeks (2 weeks imbibed followed by 8 weeks at controlled moisture content) or more, significantly improved the performance at the germination temperatures 15 and 30°C. For Pseudotsuga menziesii seed pretreatment at controlled moisture content for up to 2 + 12 weeks was superior, especially at low germination temperatures, and it was possible to redry pretreated seed to 8% moisture content without loss of the pretreatment effect or germination capacity. For Picea sitchensis seeds the controlled moisture content method for a duration of 7-10 weeks proved efficient and pretreated seed of this species also tolerated redrying. It is recommended that these improved pretreatment methods should be introduced into the nurseries.

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36.
Ross, S.D. and R.C. Bower. 1989. Cost-effective promotion of flowering in a Douglas-fir seed orchard by girdling and pulsed stem injection of gibberellin A4/7. Silvae-Genetica 38(5-6): 189-195.

Keywords:      seed orchard management

                        reproduction

                        economics

Abstract: In a seed orchard in British Columbia, Canada, Douglas fir (Pseudotsuga menziesii) grafts of 5-14 cm diam. received different combinations of partial saw-cut stem girdles and ethanolic solutions of GA4/7 injected into shallow holes drilled around the main stem. Grafts averaged 79 seed-cone buds and 4500 pollen-cone buds each without treatment. Girdling alone increased production per tree of seed- and pollen-cone buds to 325 and 9300 respectively. GA4/7 alone was nearly as effective as girdling alone, the response being marginally greater at the high than low dosage (3.82 vs. 1.27 mg/cmsuperscript 2 of stem cross sectional area), but independent of whether the total dose was applied all at once or over two or three injections at 2-wk intervals. Together, girdling and GA4/7 had an additive effect on flowering, increasing production per tree of seed- and pollen-cone buds to 585 and 18 250. The combined treatment was particularly effective on smaller trees that flowered poorly or not at all without treatment, while also enhancing production significantly on larger trees. The combined treatment was safe and highly cost effective. It cost $63.75 per year to maintain each tree in the orchard, so that without any treatment the cost per seed-cone bud initiated was $0.91. Girdling (at $2.07 per tree) reduced this cost to $0.20, and girdling + GA4/7 (at $7.87 per tree) to only $0.13.

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37.
Ross, S.D.