1. Abdel-Gadir, A.Y., R.L. Krahmer and M.D. McKimmy. 1993. Relationships between intra-ring variables in mature Douglas-fir trees from provenance plantations. Wood-and-Fiber-Science 25(2): 182-191.

Keywords:      genetic tree improvement

                        wood quality

                        genetic relationships

Abstract: Relationships among a variety of densitometric characteristics of juvenile and mature wood from 360 trees growing in two plantations of Douglas fir (Pseudotsuga menziesii) in Oregon and Washington were examined. Variables included earlywood density (EWD) and width (EWW); latewood density (LWD), width (LWW), and proportion (LWP); average ring density (RD); and total ring width (RW). The RD components (EWD and LWD) had strong phenotypic and genetic correlations with their respective RW components (EWW and LWW). However, no phenotypic correlation existed between average RD and total RW, and genotypic correlation was weak. The relation between wood density and radial growth varied by plantation and genotype. It was shown that the potential exists for improving wood density in juvenile and mature wood by selection, with only a minor effect on radial growth. Selection during the juvenile period to improve mature wood quality would be feasible for RD, EWD, LWW, and LWP. Further, selection to improve juvenile RW would not result in reduced wood density during maturity.

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2.
Adams, W.T., S.N. Aitken, D.G. Joyce, G.T. Howe and J. Vargas-Hernandez. 2001. Evaluating efficacy of early testing for stem growth in coastal Douglas-fir. Silvae-Genetica 50(3/4): 167-175.

Keywords:      genetic tree improvement

                        nursery operations

                        growth

                        genetic relationships

Abstract: In a test to evaluate the ability to predict stem growth of families in the field from nursery performance (i.e., early testing), 67 open-pollinated families and 66 full-sib families of coastal Douglas-fir (Pseudotsuga menziesii var. menziesii) were sown in two nursery conditions, each replicated as separate experiments: two bareroot nursery trials established in successive years in the same nursery, and two container-sown greenhouse trials sown in different greenhouses in the same year. First year heights in the seedling trials were compared to mean stem volumes of the same open-pollinated families in eight 15-year-old field progeny tests and the same full-sib families in eleven 12-year-old tests. Family mean nursery-field correlations (rxy) were similar for all four seedling trials for both open-pollinated (OP) and full-sib (FS) families, and generally ranged between 0.30 and 0.40. Although low, it is shown that nursery-field correlations of this magnitude can be quite useful in tree improvement programmes. For example, based on the data in this study, it is estimated that a single stage of family selection for first year seedling height would be about 50% as effective in improving 15-year volume as direct selection for this trait in field tests. Early testing, however, is probably of more practical significance as a tool for culling families prior to out-planting field tests in two-stage selection schemes. It is estimated that 25% of the OP families in this study could have been culled in an early test (first stage selection), with gain in 15-year volume after subsequent field testing and selection of the remaining families (second stage selection) being nearly the same as if all families had been field tested. Thus, early testing is an effective tool for reducing the size and cost of field progeny tests without sacrificing genetic gain.

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3.
Adams, W.T., J. Zuo, J.Y. Shimizu, J.C. Tappeiner and J.H. Zuo. 1998. Impact of alternative regeneration methods on genetic diversity in coastal Douglas-fir. Forest Science 44:390-396.

Keywords:      planting operations

                        genetic relationships

Abstract: Genetic markers (17 allozyme loci) were used to study the genetic implications of natural and artificial regeneration following 3 regeneration methods (group selection, shelterwood, and clearcut) in coastal Douglas fir (Pseudotsuga menziesii var. menziesii) forests in the Willamette Valley, Oregon, USA. In general, harvesting followed by either natural or artificial regeneration resulted in offspring populations which were very similar to those in the previous generation. The removal of the smallest trees to form shelterwoods resulted in the removal of rare, presumably deleterious, alleles. Fewer alleles per locus were observed among residual trees (2.76) and natural regeneration (2.75) than were found in uncut (control) stands (2.86). The shelterwood regime appeared quite compatible with gene conservation, and it was considered that it was best to leave parent trees of a range of sizes to act as gene conservation reserves, in order to maximize the number of alleles (regardless of current adaptive value) in naturally regenerated offspring. Seedling stocks used for artificial regeneration in clearcut, shelterwood, and group selection stands (7 total) had significantly greater levels of genetic diversity, on average, than natural regenerated seedlings.

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4.
Aitken, S.N. and W.T. Adams. 1996. Genetics of fall and winter cold hardiness of coastal Douglas-fir in Oregon. Canadian-Journal-of-Forest-Research 26(10): 1828-1837.

Keywords:      genetic tree improvement

                        tree/stand protection

                        tree/stand health

                        tree physiology

                        genetic relationships

Abstract: Genetic variation in autumn cold hardiness was studied in two western Oregon breeding populations of coastal Douglas fir (Pseudotsuga menziesii var. menziesii), one on the west slope of the Cascade Mountains and the other in the Coastal Range. On six sampling dates (September, October and November 1992 and January, September and October 1993), shoot cuttings from 40 open-pollinated families in each of two progeny test sites for each breeding zone were subject to artificial freezing at two test temperatures. Damage in each shoot was recorded as visible injury to needle, stem and bud tissue separately. Considerable family variation was found for cold injury scores in all tissues in early to mid autumn, but differences were often smaller or nonsignificant in late autumn and midwinter. Individual heritability estimates for needle cold injury were low (<0.40) and generally decreased in late autumn and midwinter. Family rankings for autumn cold hardiness, however, are expected to be relatively consistent over sites and years, although needles appear to display more family-by-site interaction than stems or buds. Genetic correlations between tissues in cold injury varied considerably and were sometimes weak, indicating that the evaluation of a single tissue is probably not adequate for assessing overall cold hardiness of genotypes. Autumn and winter cold hardiness seem to be largely under separate genetic control since genetic correlations between hardiness at these two stages were weak. This study confirms earlier results in Washington breeding populations and shows that coastal Douglas fir families can be effectively ranked for autumn cold hardiness by conducting artificial freeze tests on cut shoots in mid-autumn (October) and scoring damage to stems and at least one other tissue.

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5.
Aitken, S.N. and W.T. Adams. 1997. Spring cold hardiness under strong genetic control in Oregon populations of Pseudotsuga menziesii var. menziesii. Canadian-Journal-of-Forest-Research 27(11): 1773-1780.

Keywords:      genetic tree improvement

                        tree/stand protection

                        tree/stand health

                        tree phenology

                        tree physiology

                        genetic relationships

Abstract: Genetic variation in spring cold hardiness of shoots prior to bud break was studied in two Oregon breeding populations of Pseudotsuga menziesii var. menziesii, one on the west slope of the Cascade Mountains and the other in the Coast Range. In March and April 1993, and April 1994, shoot cuttings from 40 open-pollinated families in each of two progeny test sites in each breeding zone were subjected to artificial freezing. Visible cold damage to needle, stem, and bud tissues was recorded. Date of bud burst (all sites), and injury resulting from a 1992 natural frost event (one site), were also recorded. Spring cold injury varied widely among families. Individual heritabilities for spring cold injury scores averaged 0.76 in the Coastal zone and 0.42 in the Cascade zone. Genetic correlations among tissues, sites, sampling dates, and years, and between April cold injury and date of bud burst were high, in most cases over 0.80. Correlations were also strong between natural frost damage in 1992 and artificial cold injury scores in 1993. Artificial freeze testing stem tissues of cut shoots sampled in April from a single test site should effectively rank families in this region for spring cold hardiness.

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6.
Aitken, S.N., W.T. Adams, N. Schermann and L.H. Fuchigami. 1996. Family variation for fall cold hardiness in two Washington populations of coastal Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco). Forest-Ecology-and-Management 80(1/3): 187-195.

Keywords:      genetic tree improvement

                        tree/stand protection

                        tree/stand health

                        tree physiology

                        tree phenology

                        genetic relationships

Abstract: In order to assess the genetics of autumn (fall) cold hardiness in coastal Douglas fir (Pseudotsuga menziesii var. menziesii), shoot cuttings were collected in October from saplings (9-year-old trees) of open-pollinated families in two progeny tests in each of two breeding zones in Washington, one in the Coast range (80 families) and one on the west slope of the Cascade Mountains (89 families). Samples from over 5500 trees were subjected to artificial freezing and visually evaluated for needle, stem and bud tissue injury. The extent to which cold injury is genetically related to tree height and shoot phenology (timing of bud burst and bud set) was also evaluated. Significant family variation was found for all cold hardiness traits; however, individual heritability estimates were relatively low (ranging from 0.09 to 0.22). Significant family-by-test site interaction was detected for needle injury in the Cascade breeding zone, but not in the coastal zone. Genetic correlations (rA) among needle, stem and bud tissues for cold damage were weak (0.16<less or =>rA<less or =>0.58) indicating that genes controlling autumn cold hardening are somewhat different for different tissues. Timing of bud burst and bud set were only weakly correlated with cold injury (rA<less or =>0.49). Thus, bud phenology is a poor predictor of autumn cold hardiness in this species. There was no consistent relationship between tree height and cold injury in the coastal zone. In the Cascade zone, taller trees appeared to be more susceptible to cold injury, but the association was weak (mean rA=0.38, range 0.20-0.72).

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7.
Anekonda, T.S., M.C. Lomas, W.T. Adams, K.L. Kavanagh and S.N. Aitken. 2002. Genetic variation in drought hardiness of coastal Douglas-fir seedlings from British Columbia. Canadian-Journal-of-Forest-Research 32(10): 1701-1716.

Keywords:      genetic tree improvement

                        tree/stand protection

                        growth

                        tree/stand health

                        genetic relationships

                        tree physiology

Abstract: Genetic variation in drought hardiness traits and their genetic correlations with growth potential and recovery traits were investigated in 39 full-sib families of coastal Douglas-fir (Pseudotsuga menziesii var. menziesii) from southwestern British Columbia, Canada. Seedlings of these families were grown in raised nursery beds and subjected to three moisture regimes each in the second (well-watered or control, mild, and moderate drought) and third (control, severe drought, and recovery from second-year moderate drought) seasons. Traits assessed included drought hardiness (foliage damage, cavitation of xylem tracheids, xylem hydraulic conductivity, and height and diameter growth increment) in the drought treatments, growth potential (total height and diameter) in the control treatment, and height and diameter growth increments in the recovery treatment. Xylem cavitation in the growth ring produced in a particular year was nearly three times greater under the moderate drought and four times greater under the severe drought than in the control treatment. Xylem hydraulic conductivity of seedlings in the severe drought treatment was 40% lower than conductivity of seedlings under the control treatment. Mean foliage damage in seedlings subjected to severe drought (third season) was much greater (33%) than in seedlings subjected to mild or moderate drought (second season). Families differed significantly in most drought hardiness traits, with individual tree heritabilities averaging 0.19. Thus, much potential exists for identifying drought-hardy families at the seedling stage and using this information for deployment or breeding purposes. In addition, most hardiness traits were strongly intercorrelated (genetic correlations often exceeded |0.80|) indicating that these traits are controlled largely by the same set of genes and that selection for hardiness based on one trait will increase hardiness as reflected in the other traits as well. Genetic correlations were only moderate (0.49) between hardiness traits measured in different years, perhaps due to the large difference in severity of the drought applied in the two seasons. Although injury to seedlings, as reflected in foliage damage and xylem cavitation, was relatively low under the moderate drought of the second season, it did result in reduced growth increment the following (recovery) year. Growth potential under favourable moisture regimes was nearly uncorrelated with drought hardiness, suggesting that drought hardiness could be improved in this southwestern British Columbia breeding population without negatively impacting growth potential in favourable moisture conditions.

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8.
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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9.
Birot, Y. and C. Christophe. 1983. Genetic structures and expected genetic gains from multitrait selection in wild populations of Douglas fir and Sitka spruce. I. Genetic variation between and within populations. Silvae-Genetica 32(5/6): 141-151.

Keywords:      genetic tree improvement

                        genetic relationships

                        growth

                        tree phenology

                        tree morphology

Abstract: For Douglas fir, 371 open-pollinated progenies from 26 provenances ranging from N. to S. along the western foothills of the Cascade Mts., Washington, were tested. For Sitka spruce, 292 open-pollinated progenies from 21 provenances ranging from S. British Columbia to middle-Oregon were tested. Observations were made on growth, phenology and form from the nursery stage up to age 12. Classical patterns of geographic variation were observed for both species. Heritability and genetic correlations varied from one provenance to another, especially for Douglas fir, and also changed over time. Sitka spruce showed high additive effects, offering good prospects of future genetic gains. It was concluded that preliminary investigations on genetic parameters were necessary before setting up a breeding strategy.

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10.
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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11.
Copes, D.L. 1999. Breeding graft-compatible Douglas-fir rootstocks (Pseudotsuga menziesii (MIRB.) FRANCO). Silvae-Genetica 48(3/4): 188-193.

Keywords:      genetic tree improvement

                        tree grafting

                        tree/stand protection

                        growth

                        tree phenology

                        tree/stand health

                        genetic relationships

Abstract: A study encompassing 24 years was conducted to determine if a breeding programme could produce highly graft-compatible rootstocks for P. menziesii. A total of 27 trees of apparent high graft compatibility were selected and crossed to produce 226 control-pollinated families. Seedlings were grown, field planted and grafted with test scions. Graft unions from field tests were evaluated anatomically for internal symptoms of incompatibility. Average compatibility of progeny from the 226 crosses was 90.6%, compared with 65% in native populations. Breeding values were calculated for each parent by the best linear prediction (BLP) procedure. Average compatibility resulting from crossing among the top 10 parents was estimated by breeding values as 95.4%. Field-test results of progeny from 34 crosses among the 10 most compatible parents showed 96% compatibility. In addition to field-tests for graft compatibility, nursery tests of seedlings from 124 crosses were evaluated for second-year vegetative bud flush and seedling height. It was possible, while maintaining adequately high levels of graft compatibility, to breed both for resistance to spring frost damage and for increased seedling height.

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12.
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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13.
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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14. 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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15. El Kassaby, Y.A. and A. Benowicz.
2000. Effects of commercial thinning on genetic, plant species and structural diversity in second growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) stands. Forest-Genetics 7(3): 193-203.

Keywords:      thinning

                        commercial thinning

                        genetic relationships

Abstract: The impact of commercial thinning on biodiversity was studied in two Douglas-fir (Pseudotsuga menziesii) plantations, Weeks Lake (WL) and Fairservice Creek (FC) located on southern Vancouver Island, British Columbia, Canada. The age of the stands at thinning was 53 and 70 years for FC and WL, respectively. Other tree species were also present in both sites due to natural regeneration. Biodiversity was evaluated before and after commercial thinning with respect to tree species composition/abundance, tree species genetic diversity assessed by allozyme analysis and stand structural diversity represented by the diameter class (5-cm) distribution. In addition, understorey plant species diversity was monitored in WL and FC for 4 and 5 years following thinning, respectively. Tree species composition changed in both plantations in a similar fashion as the proportion of Douglas-fir increased at the expense of all other tree species. Stand structural diversity was simplified due to the decreased number of trees in small diameter classes. These results were expected since the commercial thinning was conducted to promote the growth of Douglas-fir. Genetic diversity parameters (average number of alleles per locus, percent polymorphic loci and expected heterozygosities) did not differ significantly before and after thinning; however, thinning resulted in a loss of 8 and 7 alleles across species for FC and WL, respectively. Most of the allelic loss occurred in the naturally regenerated species (93%). This allelic loss represents 7 and 6% of the total alleles present in FC and WL, respectively. Thus, only one allele was lost from the crop tree in the FC plantation. Understorey vegetation species richness decreased the year following commercial thinning and then consistently increased over time in both plantations. A total of 17 and 9 new species colonized FC and WL, respectively. One species was replaced in each plantation. In FC, diversity of the understorey plant community based on the Shannon diversity index (H) changed in an increasing linear trend that corresponded to the increase in species richness. On the other hand, H in WL remained stable. Species evenness (H/Hmax) did not change in WL and FC over the course of study except for seasonal fluctuations. Rare species diversity increased over time in both plantations.

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16. 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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17. 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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18. El Kassaby, Y.A. and R. Davidson.
1990. Impact of crop management practices on the seed crop genetic quality in a Douglas-fir seed orchard. Silvae-Genetica 39(5-6): 230-237.

Keywords:      genetic tree improvement

                        seed orchard management

                        genetic relationships

Abstract: The impact of 2 crop management practices, supplemental-mass-pollination (SMP) and overhead cooling, on levels of pollen contamination and outcrossing was assessed in a 13-year-old seedling Douglas fir (Pseudotsuga menziesii) seed orchard with the aid of 6 allozyme loci. A 2 x 2 factorial arrangement of SMP/no SMP and cooling/no cooling was applied to 4 genetically similar blocks of trees. The 4 treatment combinations used were spatially and temporally isolated by buffer blocks and 9-day reproductive bud phenology delay, respectively. The study led to the following conclusions: (1) no contamination was observed when cooling and/or SMP was used, (2) no inbreeding was observed when cooling and SMP were used concurrently and (3) SMP was effective in reducing consanguineous mating, but not to the level achieved by cooling and SMP combined. In addition, the results obtained from the control block (i.e. no cooling and no SMP) indicated that (1) isolation zones are effective in reducing contamination, and (2) randomization of trees within seed orchard blocks is associated with a high outcrossing rate.

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19. El Kassaby, Y.A. and
Y.S. Park. 1993. Genetic variation and correlation in growth, biomass, and phenology of Douglas-fir diallel progeny at different spacings. Silvae-Genetica 42(6): 289-297.

Keywords:      genetic tree improvement

                        nursery operations

                        genetic relationships

                        growth

                        carbon allocation

                        tree phenology

Abstract: Parents of coastal Douglas fir (Pseudotsuga menziesii) selected from natural stands on sites ranging from 0 to 450 m altitude on Vancouver Island and in southeastern British Columbia were crossed and the resulting 104 full-sib families evaluated for 3 years after germination. The full-sib families were produced by a disconnected diallel mating scheme, consisting of 7 sets of 6-parent partial diallels, grown under 2 spacing treatments in a nursery. The objectives of the study were to determine the extent of genetic control of growth traits, biomass distribution and allocation strategies, and vegetative phenology. Spacing had a significant effect on 6 of the 11 traits studied. Significant GCA variance was found for all traits except 1-year height. Individual tree narrow-sense heritability varied from 0.06 to 0.69 for root dry weight and vegetative phenology, respectively. Spacing x family interaction variance was significant for only 2 traits. Two harvest indices, based on total and above-ground dry weights, were used to assess dry matter allocation strategy and to explore potential usefulness in tree breeding. Both indices had similar heritability estimates and their genetic correlation was high (0.91), indicating that use of an index based on above ground dry weight is a good surrogate for that based on total dry weight. Genetic correlations among growth and biomass traits were generally high, while those correlations with the harvest indices were variable.

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20. 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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21. 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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22. 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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23.
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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24.
Jermstad, K.D., D.L. Bassoni, N.C. Wheeler, T.S. Anekonda, S.N. Aitken, W.T. Adams and D.B. Neale. 2001b. Mapping of quantitative trait loci controlling adaptive traits in coastal Douglas-fir. II. Spring and fall cold-hardiness. Theoretical-and-Applied-Genetics 102(8): 1152-1158.

Keywords:      genetic tree improvement

                        tree/stand protection

                        genetic relationships

                        tree physiology

                        tree phenology

Abstract: Quantitative trait loci (QTLs) affecting fall and spring cold hardiness (cold resistance) were identified in a three-generation outbred pedigree of coastal Douglas-fir (Pseudotsuga meniziesii var. menziesii [Pseudotsuga menziesii]) in a field experiment conducted in Washington and Oregon, USA, during 1996-97. Eleven QTLs controlling fall cold hardiness were detected on four linkage groups, and 15 QTLs controlling spring cold hardiness were detected on four linkage groups. Only one linkage group contained QTLs for both spring and fall cold hardiness, and these QTLs tended to map in close proximity to one another. Several QTLs were associated with hardiness in all three shoot tissues assayed in the spring, supporting previous reports that there is synchronization of plant tissues during de-acclimatization. For fall cold hardiness, co-location of QTLs was not observed for the different tissues assayed, which was consistent with previous reports of less synchronization of hardening in the fall. In several cases, QTLs for spring or fall cold hardiness mapped to the same location as QTLs controlling spring bud flush. QTL estimations, relative magnitudes of heritabilities, and genetic correlations based on clonal data in this single full-sib family, supports conclusions about the genetic control and relationships among cold hardiness traits observed in population samples of Douglas-fir in previous studies.

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25.
Johnson, G.R. 1997. Site-to-site genetic correlations and their implications on breeding zone size and optimum number of progeny test sites for coastal Douglas-fir. Silvae-Genetica 46(5): 280-285.

Keywords:      genetic tree improvement

                        genetic relationships

Abstract: Type B genetic correlations were used to examine the relationship among geographic differences between sites and their site-to-site genetic (Type B) correlations. Examination of six local breeding zones in Oregon indicated that breeding zones were, for the most part, not too large because few environmental variables were correlated with Type B genetic correlations. The data also were used to examine expected gains from using combinations of sites in selection indices. Even though additional sites always increased the expected genetic gain, the marginal increase was only minimal if 3 or 4 sites were already in the index. The trend was consistent over all 6 breeding zones.

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26.
Johnson, G.R. 2002. Genetic variation in tolerance of Douglas fir to Swiss needle cast as assessed by symptom expression. Silvae-Genetica 51(2/3): 80-86.

Keywords:      genetic tree improvement

                        tree/stand protection

                        tree/stand health

                        growth

                        genetic relationships

Abstract: The incidence of Swiss needle cast (caused by Phaeocryptopus gaeumannii) on Douglas fir (Pseudotsuga menziesii) has increased significantly in recent years on the Oregon coast. Genetic variation in symptoms of disease infection, as measured by foliage traits, was assessed in two series of progeny trials to determine whether these "crown health" indicators were under genetic control and correlated with tolerance (tolerance being continued growth in the presence of high disease pressure). Foliage traits generally had lower heritabilities than growth traits and were usually correlated with diameter growth. Foliage traits of crown density and colour appeared to be reasonable indicators of disease tolerance. In the absence of basal area data, assessing crown density and colour can help screen for families that show tolerance to the disease.

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27.
Johnson, G.R., R.A. Sniezko and N.L. Mandel. 1997. Age trends in Douglas-fir genetic parameters and implications for optimum selection age. Silvae-Genetica 46(6): 349-358.

Keywords:      genetic tree improvement

                        growth

                        genetic relationships

Abstract: Trends in genetic variation in Douglas fir (Pseudotsuga menziesii) were examined over 51 progeny test sites throughout western Oregon. Narrow sense heritabilities for height and diameter showed an increasing trend to age 25, the oldest age examined. Before age 10, height heritabilities were relatively unstable. Type B site-site genetic correlations increased slowly with age for height and remained relatively stable for diameter. Age-age correlations were used to develop an equation to predict age-age correlations by using the log of the age ratios (LAR). Optimum selection age was calculated for a 60-year rotation by using two measures of efficiency: gain per year and discounted gain. The optimum selection age for height tended to be 2 to 3 years earlier than for diameter. Gain per year was maximized at age 10 for height and age 13 for diameter.

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28.
Johnson, R. 1998. Breeding design considerations for coastal Douglas-fir. Pacific-Northwest-Research-Station, USDA-Forest-Service General-Technical-Report PNW-GTR-411. iii + 34 p.

Keywords:      genetic tree improvement

                        seed orchard management

                        genetic relationships

Abstract: The basic principles of designing forest tree breeding programmes are reviewed for Douglas fir (Pseudotsuga menziesii) in the Pacific Northwest (USA). Breeding populations are discussed given current and future breeding zone sizes and seed orchard designs. Seed orchard composition is discussed for potential genetic gain and maintaining genetic diversity in the forest. Mating and field testing designs are described and compared. Recommendations of the Breeding Zone Evaluation and Restructuring Cooperatives Working Group of the Northwest Tree Improvement Cooperative are presented.

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29.
Kaya, Z. 1992. The effects of test environments on estimation of genetic parameters for seedling traits in 2-year-old Douglas-fir. Scandinavian-Journal-of-Forest-Research 7(3): 287-296.

Keywords:      genetic tree improvement

                        nursery operations

                        genetic relationships

                        growth

                        tree phenology

Abstract: The effects of test environments (dry versus wet) on the estimation of genetic parameters in seedling traits were studied in 160 open-pollinated families of Douglas fir (Pseudotsuga menziesii) from southwestern Oregon. Seedlings from four populations were grown in two test nursery environments between which a water potential difference of -9 bars was created over two growing seasons, by withholding water for 4 and 8 wk, respectively. Estimated genetic variances in most growth and phenology traits were considerably higher for seedlings grown in the wet environment than for those in the dry. Estimated genetic correlations between the same traits measured in different test environments indicated that most seedling traits studied for two growing seasons were genetically stable in both environments, suggesting that genotype environment interaction in these traits are weak. However, it is emphasized that the effect of test environment on estimation of genetic parameters in seedling traits, especially in adaptive seedling traits, should be evaluated very carefully when early evaluation of genetic entries is practised in Douglas-fir, since these traits (budburst timing, lammas growth and free growth) appear to be plastic in character.

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30.
Kaya, Z. 1993. Genetic variation in shoot growth components and their correlations in Pseudotsuga menziesii var. menziesii seedlings. Scandinavian-Journal-of-Forest-Research 8(1): 1-7.

Keywords:      genetic tree improvement

                        genetic relationships

                        growth

Abstract: Stem-unit measurements could be useful for early selection if these dimensions were highly heritable and strongly correlated with traits of commercial interest, such as height growth. Height increments and the number and length of stem segments were measured in the first and second growth period in predetermined and free growth of Douglas-fir (Pseudotsuga menziesii var. menziesii) seedlings from 40 families in each of 2 populations from southwestern Oregon, USA. Populations, or families in populations, exhibited genetic variation in all traits except number of stem units in predetermined growth. Heritabilities for stem-unit measurements were higher than those for height increment in the first growing season, but not in the second. Correlations among measurements of stem units and height increments were only moderate (generally <0.70). It is concluded that stem units are not likely to be better measurements of height in early selection of Douglas-fir than are measurements of either predetermined or free growth.

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31.
Kaya, Z., R.K. Campbell and W.T. Adams. 1989. Correlated responses of height increment and components of increment in 2-year-old Douglas fir. Canadian-Journal-of-Forest-Research 19(9): 1124-1130.

Keywords:      genetic tree improvement

                        tree phenology

                        growth

                        genetic relationships

Abstract: The consequences for growth and phenology of early selection for height or its growth components were evaluated in 160 open-pollinated families of Douglas fir (Pseudotsuga menziesii) from SW Oregon. Seedlings from 2 inland and 2 coastal populations (40 families each) were grown for 2 growing seasons in a 'moist' and 'dry' nursery environment. Predicted response to selection suggests that risk of low juvenile-mature correlation and maladaptation with early selection would be less in the inland than in the coastal region. Early bud set in the 1st yr was genetically correlated with larger overwintering buds in seedlings from both inland and coastal regions. These larger buds yielded a large increment of predetermined growth in the 2nd yr, followed by little or no free growth and early bud set. Seedlings with late bud set in the 1st yr had the converse pattern. Inland seedlings set buds much earlier on av. than coastal seedlings, hence seedlings from the 2 regions had different growth patterns. Risks that can attend early selection for ht. generally would be decreased in both regions by selecting for predetermined growth, but several qualifications are discussed.

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32.
Kimball, B.A., G.R. Johnson, D.L. Nolte and D.L. Griffin. 1999. An examination of the genetic control of Douglas-fir vascular tissue phytochemicals: implications for black bear foraging. Forest-Ecology-and-Management 123(2/3): 245-251.

Keywords:      genetic tree improvement

                        tree/stand</