1. 1994. Annual Report - Forest Research Laboratory, Oregon State University. Forest Research Laboratory, Oregon State University. ii + 29 pp.

Keywords: genetic tree improvement

tree/stand protection

tree phenology

tree physiology

Abstract: Highlights of research conducted during 1993-1994 are presented, including: preliminary results of a 2-year (1992-94) field cold hardiness study of Douglas fir [Pseudotsuga menziesii] in Oregon, USA involving open pollinated progeny of 40 parents at a high and a low elevation; results of a progeny test on the frequency of second flushing of Douglas fir near Orleans, France; and variation in stable carbon isotope ratios (a measure of water use efficiency) among varieties and populations (coastal and Rocky Mountain) of Douglas fir.

2. 1997. PNWTIRC Annual Report 1996-97, Pacific Northwest Tree Improvement Research Cooperative. Oregon State University, Oregon, USA. ii + 29 p.

Keywords: genetic tree improvement

tree/stand protection

tree phenology

growth

wood quality

tree physiology

Abstract: The report describes highlights for 1996-97, current research (3 projects), student project updates (3 projects), planned Douglas fir [Pseudotsuga menziesii] seed orchards for the new millennium and other planned activities of the PNWTIRC, a research cooperative operating in the Pacific Northwest area of North America (USA and Canada). Details are included of publications and finances. Details of the 3 current research projects and the 3 student projects, which all concern Douglas fir, are presented as short papers including brief results: (1) Influence of second flushing on cold hardiness; (2) Seedling drought physiology study; and (3) Quantitative trait loci influencing cold hardiness; (4) Seedling cold hardiness; (5) Growth response of saplings to drought; and (6) Measurement study follow-up: age-age correlations in forking defects.

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3. 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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4. 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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5. 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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6. Blake, J.I., R. Linderman and D. Lavender. 1988b. Seedling vigor of Douglas fir and western hemlock in relation to ethylene exposure levels and ethane production during cold storage. 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. 235-242.

Keywords: nursery operations

tree morphology

tree phenology

tree/stand health

Abstract: The effects were examined of ethylene treatment on Pseudotsuga menziesii and Tsuga heterophylla 2-yr-old bare root seedlings lifted in October or December (in a nursery in Washington) and stored for 2 and 7 days. Seedlings exposed to 0.5 p.p.m. ethylene at +1 degrees C for 7 days exhibited reduced bud and root activity. These effects were apparently reversed at 5.0 p.p.m. In a second experiment, reducing ethylene concentrations with KMnO4 during storage at +1 or +10 degrees C for 30 days did not affect foliage colour, but root and bud activity were generally enhanced. Ethylene concentrations in control bags ranged from 0.80 to 2.24 p.p.m. in October-lifted seedling bags and from 0.10 to 1.3 p.p.m. in December-lifted samples. The quantity of ethane in stored bags of P. menziesii seedlings was closely related to an increase in foliage discoloration. Little or no ethane was measured in T. heterophylla storage bags.

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7. Blake, J.I. and R.G. Linderman. 1992. A note on root development, bud activity, and survival of Douglas-fir, and survival of western hemlock and noble-fir seedlings, following exposure to ethylene during cold storage. Canadian-Journal-of-Forest-Research 22(8): 1195-1200.

Keywords: nursery operations

tree morphology

tree phenology

tree/stand health

Abstract: Three cold storage experiments were conducted with bare-root (2+0) Douglas fir (Pseudotsuga menziesii) seedlings from coastal Oregon and eastern Washington Cascade sources. The objectives were to determine the effects of ethylene at ambient and below-ambient (absorbed by KMnO4 pellets) concentrations, and at 0.5 and 5 p.p.m. ethylene, during short-term storage on subsequent root development and bud activity, and to relate these results to survival in the field at sites in Washington and Oregon, after prolonged cold storage. Root numbers and lengths were measured 28 days after a 7-day storage period after lifting seedlings on 27 September and 1 December. In the coastal source, root numbers and lengths in the 5 p.p.m. ethylene treatment were, respectively, 46 and 49% greater in September, and 22 and 13% greater in December, than the controls. No comparable treatment effects were found for the Cascade source. Neither the KMnO4 nor the 0.5 p.p.m. ethylene treatments affected root development in either seed source. For terminal buds in the controls, the number of days to 50% bud break was increased 2-8 days by a 30-day cold storage period compared with a 7-day period. For the coastal source, no increase in the time to 50% bud break was observed in the 5 p.p.m. ethylene treatment. Seedling survival was evaluated in the field for the same treatments following 4 months cold storage for the Douglas fir sources, coastal western hemlock (Tsuga heterophylla), and noble fir (Abies procera). Survival for the 5 p.p.m. ethylene treatment compared with the control was increased by 55% in the coastal Douglas fir source and by 13% in western hemlock. These results suggest that stimulated root development and bud activity may be partially responsible for the observed survival increase following cold storage at elevated ethylene levels.

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8. Childs, S.W. and L.E. Flint. 1987. Effect of shadecards, shelterwoods, and clearcuts on temperature and moisture environments. Forest-Ecology-and-Management 18(3): 205-217.

Keywords: planting operations

tree/stand health

soil properties

tree physiology

tree phenology

Abstract: A comparison was made of two common techniques used to improve seedling survival on hot, dry reforestation sites. Adjacent shelterwood and clearcut sites in SW Oregon, USA, planted with 2+0 Douglas fir, were located and instrumented to compare temp. and moisture. In addition, cardboard shadecards were placed beside half of the seedlings studied. Seasonal measurements or observations of soil moisture, soil temp., solar radiation, air temp., stomatal diffusion resistance, seedling phenology and survival provided the basis for comparisons. Shelterwoods and shadecards improved seedling survival in relation to the clearcut. Both treatments affected soil temp. but the nature of the effects was different. The shelterwood canopy reduced solar radiation incident at the soil surface and caused cooler soil temp. throughout the soil profile. Shadecards reduced soil temp. only to a depth of 20 mm. Both treatments reduced the duration of periods of high soil temp. Shelterwood treatment delayed seasonal water loss and reduced seedling water stress as measured by stomatal resistance. Shadecards did not significantly affect seedling stomatal resistance. Differences in seedling survival caused by shadecards and shelterwoods are apparently due to different influences on the seedling microclimate. Shelterwood causes a large reduction in soil temp. as well as decreased seedling water stress. Shadecards modify the soil temp. less extensively and so have less effect on seedling survival.

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9. Copes, D.L. 1983. Failure of grafted Douglas-fir planted at Monterey, Calif. Tree-Planters' Notes 34(3): 9-10.

Keywords: genetic tree improvement

tree grafting

growth

tree/stand health

tree phenology

Abstract: Trees were grafted onto established rootstocks (grown from rooted cuttings from Oregon or seedlings from California) in Oregon in April 1979, and in Dec. 109 of these were transplanted to a site on the Monterey Peninsula, California. Grafts were examined in July 1981. More than 30% of the transplanted grafts had died of early incompatibility and the cumulative 1980 and 1981 leader growth of the survivors averaged 12 cm. Identical grafts in Oregon showed 2-10% death due to incompatibility and av. leader growth of 1-2 m. It is suggested that the atypical unreliable budflush and reduced leader elongation in Monterey was due to winter temperatures that were not cool enough to satisfy bud dormancy requirements. It had been hoped to establish seed orchards out of range of pollen contamination from local Douglas fir stands.

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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. Dunsworth, B.G. 1988. Douglas-fir fall root phenology: a bioassay approach to defining planting windows. 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. 295-307.

Keywords: planting operations

growth

tree/stand health

tree phenology

Abstract: Natural seedling root phenology during the autumn and spring was observed in Douglas fir [Pseudotsuga menziesii] and western hemlock [Tsuga heterophylla] over several seasons on Vancouver Island, British Columbia. These observations were used to determine the soil climate conditions under which peak activity occurred. Mitotic indexing and total root tip counts were used to quantify root activity. Peak activity for both spring and autumn in both species could be reasonably bracketed by soil climate conditions of -1 bar soil tension and 4 degrees C. The hypothetical planting window defined by these soil climate conditions was then tested with a series of timing of planting studies for Douglas fir (autumn) and western hemlock (spring) containerized stock. The results indicated that although quality of planting stock and the season following planting are influential, a 10 to 15% survival and growth advantage can be gained by planting within the hypothetical window.

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13. Duryea, M.L. and S.K. Omi. 1987. Top pruning Douglas-fir seedlings: morphology, physiology, and field performance. Canadian-Journal-of-Forest-Research 17(11): 1371-1378.

Keywords: nursery operations

nursery pruning

tree phenology

tree/stand health

growth

yield

Abstract: Seedlings from 9 seed sources at 6 nurseries in Washington, Oregon and California were treated with various pruning treatments including tall and short ht. (25 and 15 cm, respectively), early and late timing (6 wk after bud burst or 6 wk after bud set, respectively), pruning twice or no pruning. Seedlings were evaluated for phenology and quality, and graded in the nursery. For each seed source, seedlings were planted at field sites in their own zone and on one common site. Seedlings pruned tall and early began growing again within 5 wk and set buds 2 wk later than unpruned seedlings. Shippable yield of seedlings pruned tall and early and of unpruned seedlings were n.s.d, although more pruned seedlings had multiple leaders. Pruned seedlings were smaller than unpruned seedlings at the time of planting. Survival and growth were the same for pruned and unpruned seedlings in the 1st year after planting. Pruned seedlings grew more than unpruned seedlings in the 2nd year, but were still shorter after 2 yr. Field growth was greater in seedlings pruned tall or early than in seedlings pruned short or late. It is concluded that pruning should be continued as a cultural practice if it benefits nurseries, but that late short pruning should be avoided.

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14. 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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15. 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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16. 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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17. 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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18. 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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19. Folk, R.S., S.C. Grossnickle, P. Axelrood and D. Trotter. 1999. Seed lot, nursery, and bud dormancy effects on root electrolyte leakage of Douglas-fir (Pseudotsuga menziesii) seedlings. Canadian-Journal-of-Forest-Research 29(8): 1269-1281.

Keywords: nursery operations

tree physiology

tree/stand health

photosynthesis

growth

tree phenology

Abstract: The effects of seed lot, nursery culture, and seedling bud dormancy status on root electrolyte leakage (REL) of Douglas-fir (Pseudotsuga menziesii) seedlings were assessed to determine if these factors should be considered when interpreting REL for seedling quality. The relationships of REL to survival, net photosynthesis (Pn), stomatal conductance (gwv) mid-day shoot water potential ( Psi mid), root growth capacity (RGC), and relative height growth were determined for each factor in experiments in 1994-95 in nurseries in British Columbia. Nursery culture had no effect on the relationship between REL and all other measured attributes. Seed lot affected the relationship between REL and Pn, Psi mid, and survival. However, critical REL (i.e., lowest value associated with detectable root damage) and PS80 REL (i.e., level associated with an 80% probability for survival) were similar between seed lots. Bud dormancy status affected the relationship between REL and survival, RGC, and relative height growth. Control levels of REL, critical REL, and PS80 REL decreased as the number of days required for 50% terminal bud break declined. Thus, terminal bud dormancy status must be known before REL can be used to assess seedling quality. If the bud dormancy status of Douglas-fir populations is known, then critical and PS80 REL levels may be useful as indices of root damage.

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20. Jermstad, K.D., D.L. Bassoni, K.S. Jech, N.C. Wheeler and D.B. Neale. 2001a. Mapping of quantitative trait loci controlling adaptive traits in coastal Douglas-fir. I. Timing of vegetative bud flush. Theoretical-and-Applied-Genetics 102(8): 1142-1151.

Keywords: genetic tree improvement

tree phenology

Abstract: Thirty-three unique quantitative trait loci (QTLs) affecting the timing of spring bud flush have been identified in an intraspecific mapping population of coastal Douglas-fir (Pseudotsuga menziesii var. menziesii) in a field experiment conducted in several sites in Washington and Oregon, USA, during 1995-98. Both terminal and lateral bud flush were measured over a 4-year period on clonal replicates at two test sites, allowing for the repeated estimation of QTLs. QTLs were detected on 12 linkage groups and, in general, each explained a small proportion of the total phenotypic variance and were additive in effect. Several QTLs influenced the timing of bud flush over multiple years, supporting earlier evidence that the timing of bud flush through developmental stages was under moderate to strong genetic control by the same suite of genes through developmental stages. However, only a few QTLs controlling the timing of bud flush were detected at both test sites, suggesting that geographic location plays a major role in the phenology of spring growth. A small number of QTLs with year and site interactions were also estimated.

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21. 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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22. 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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23. 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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24. Khan, S.R., R. Rose, D.L. Haase and T.E. Sabin. 1996. Soil water stress: its effects on phenology, physiology, and morphology of containerized Douglas-fir seedlings. New-Forests 12(1): 19-39.

Keywords: nursery operations

growth

tree physiology

tree phenology

Abstract: Containerized 3-month-old Douglas fir (Pseudotsuga menziesii) seedlings were subjected to six moisture-stress treatment (ranging from 7 to 65% soil water content by volume) for 12 weeks. At the end of this period, there were significant differences in phenological, physiological, and morphological responses among the seedlings in the various moisture-stress treatments. In general, seedlings grown under very high or very low soil moisture conditions were adversely affected, while those grown under moderate conditions (29 to 53% water content) exhibited optimum growth, bud development, and nutrient and starch reserves. The use of vector analysis was found to be helpful in data interpretation. The results indicate the importance of closely monitoring nursery moisture regimes in order to achieve the best seedling quality.

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25. Lavender, D.P. and S.G. Stafford. 1985. Douglas-fir seedlings: some factors affecting chilling requirement, bud activity, and new foliage production. Canadian-Journal-of-Forest-Research 15(2): 309-312.

Keywords: nursery operations

tree/stand health

tree physiology

tree phenology

Abstract: Potted 2-yr-old seedlings were exposed to a range of natural and artificial environments at the Forest Research Laboratory, Oregon State University during the autumn and winter before a 9-wk period in an environment designed to permit active shoot growth. Seedling response demonstrated (i) that exposure to a period of short (9 h), mild (20 degrees C) days prepared seedlings for the beneficial effects of subsequent chilling (4.4 degrees C) temperatures and (ii) that the physiology of dormancy of Douglas-fir seedlings may be adversely affected by environments that differ markedly from those prevailing in the Pacific Northwest during the autumn.

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26. Li, P. and W.T. Adams. 1993. Genetic control of bud phenology in pole-size trees and seedlings of coastal Douglas-fir. Canadian-Journal-of-Forest-Research 23(6): 1043-1051.

Keywords: genetic tree improvement

genetic relationships

tree phenology

growth

Abstract: The extent to which bud phenology is genetically controlled and related to growth traits was examined in seedlings and pole-size trees of coastal Douglas-fir (Pseudotsuga menziesii var. menziesii). Data on bud burst, bud set, and stem growth were collected from pole-size trees of 60 open-pollinated families growing in four plantations in Oregon, and from seedlings of 45 of these same families growing in a bare-root trial in Washington, and greenhouse and transplant trials in Washington and Oregon. In both age-classes, bud burst was under moderate to strong genetic control (hsuperscript 2<more or =>0.44) and family breeding values were stable across test environments, indicating that this trait could be readily altered in breeding programmes. Bud set was inherited strongly in pole-size trees (hsuperscript 2=0.81) but weakly in seedlings (hsuperscript 2<less or =>0.30). Both bud burst and bud set were positively correlated with growth in seedlings and pole-size trees. Thus, selection for greater growth at either age-class is expected to delay bud burst and bud set. An evaluation was made of the accuracy of two alternatives for assessing bud burst phenology in pole-size trees compared with the traditional method. It was shown that bud-burst date on lateral branches can be used to rank accurately both individuals and families for bud-burst date on less accessible leader shoots. In addition, it was found that families can be ranked for mean bud-burst date by the proportion of trees per family that have flushed on a given scoring day. This method is only effective, however, when between 25 and 75% of all trees in the test have flushed at the time of scoring.

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27. Li, P. and W.T. Adams. 1994. Genetic variation in cambial phenology of coastal Douglas-fir. Canadian-Journal-of-Forest-Research 24(9): 1864-1870.

Keywords: genetic tree improvement

genetic relationships

tree phenology

growth

Abstract: The objectives of this study were to (i) determine the extent of genetic variation and genetic control of cambial phenology in coastal Douglas fir (Pseudotsuga menziesii var. menziesii), (ii) assess the degree to which cambial phenology is genetically related to bud-burst timing, (iii) examine genetic relationships between cambial phenology and growth traits, and (iv) evaluate the potential for indirectly altering cambial phenology in breeding programmes when selection is for stem volume. Dates of diameter-growth initiation and cessation, and duration of diameter growth (i.e., cambial phenology traits), as well as diameter increment for the 1987 growing season (15-yr-old), were estimated from cumulative diameter growth curves of individual trees of 60 open-pollinated families growing in one plantation in Oregon. Data on stem height and diameter at breast height (d.b.h.), and date of bud burst in 1987 were also collected. Dates of diameter-growth initiation and cessation differed significantly among families, but had lower estimated individual heritabilities (<less or =>0.23) than date of bud burst (0.87). Weak genetic correlations between date of bud burst and dates of diameter-growth initiation and cessation (range -0.09 to 0.26) indicate that timing of diameter growth cannot be reliably predicted from observations of the more easily measured bud burst. Cambial phenology traits were weakly correlated with 1987 diameter increment and moderately correlated with 15-year d.b.h. and volume. Selection of parents in this study for stem volume at age 15 and subsequent crosses among them, would be expected to lead to earlier initiation of diameter growth in the offspring, and possibly later cessation as well. The practical implications of these indirect responses in terms of increased risk of frost damage are unclear, since projected changes are small (i.e., a few days).

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28. Lopushinsky, W. and T.A. Max. 1990. Effect of soil temperature on root and shoot growth and on budburst timing in conifer seedling transplants. New-Forests 4(2): 107-124.

Keywords: planting operations

tree morphology

tree phenology

tree physiology

Abstract: Results are presented of studies of tree seedling morphological responses to a wide range (0-30 degrees C) of soil temperatures and the effects of soil temperature on needle water potential and stomatal conductance in ponderosa pine (P. ponderosa). Bare-root 2-year-old seedlings from high-altitude seed sources in the Cascade Mountains (Washington and Oregon states) were used for the morphological study. Root growth in Douglas fir (Pseudotsuga menziesii), Pacific silver fir (Abies amabilis), noble fir (Abies procera), lodgepole pine (Pinus contorta) and ponderosa pine began when soil temperature exceeded 5 degrees C. Root growth increased rapidly after 10 degrees C and attained maximum values at 20 degrees C. At 30 degrees C, no root growth occurred in the firs; in the pines, root growth was 30 to 39% of maximum. Maximum shoot growth also occurred at 20 degrees C. In ponderosa pine, height growth of seedlings from a high-altitude source was unaffected by cold soil, but in low-altitude seedlings it was reduced. Budburst in Douglas fir and the pines was delayed up to 11 days by cold soil, whereas in silver fir and noble fir, it was only slightly delayed. Prior to new root growth in ponderosa pine, xylem pressure potentials and stomatal conductances during the afternoon indicated reduced stomatal opening at all soil temperatures, whereas 23 days later, stomata were open to a greater degree when temperatures exceeded 10 degrees C. Implications of these results are briefly discussed: although root growth was initiated at soil temperatures of 5 degrees C, this does not imply that seedlings should not be transplanted until soils reach that temperature.

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29. MacDonald, J.E. 1996. Early development of bud dormancy in conifer seedlings. In Plant dormancy: physiology, biochemistry and molecular biology. Ed. G.A. Lang. Wallingford, UK: CAB INTERNATIONAL. pp. 193-199.

Keywords: nursery operations

tree phenology

Abstract: A study is reported on 1-yr-old coastal Douglas fir (Pseudotsuga menziesii var. menziesii) nursery seedlings on Vancouver Island, British Columbia, with the aim of determining whether there was a difference in dormancy development in response to short-day or moderate moisture stress treatments. Successful dormancy induction was obtained during the first week of both types of treatment, but the phenology of dormancy development was different, being earlier in the short-day treatment. Of the short-day treatments tested (3-6 wk), dormancy development was earliest for the 4-wk duration.

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30. Margolis, H.A. and R.H. Waring. 1986b. Carbon and nitrogen allocation patterns of Douglas-fir seedlings fertilized with nitrogen in autumn. II. Field performance. Canadian-Journal-of-Forest-Research 16(5): 903-909.

Keywords: nursery operations

nursery fertilization

tree phenology

growth

tree physiology

Abstract: Douglas fir seedlings (2+0), unfertilized or fertilized with ammonium nitrate in Oct. 1983, were planted out in Feb. 1984 near Corvallis, Oregon. Rye grass was grown on half the plots to induce water stress during the typical summer drought. Sucrose was applied to soil around each seedling to stimulate microbial growth and thus to immobilize nitrogen in the microbial biomass and create nitrogen stress in Douglas fir. Fertilized seedlings had earlier budbreak, produced more shoot growth and had higher relative growth rates, net assimilation rates and leaf area ratios than unfertilized seedlings. Grass significantly increased predawn moisture stress in both fertilized and unfertilized seedlings by early Aug. By 3 Sep., unfertilized seedlings growing with grass were significantly more stressed than other seedlings. Fertilizer did not have a significant effect on concn. of free amino acids and total N at the end of the growing season, but grass competition affected both N and carbohydrate chemistry.

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31. McCreary, D.D., D.P. Lavender and R.K. Hermann. 1990. Predicted global warming and Douglas-fir chilling requirements. Annales-des-Sciences-Forestieres 47(4): 325-330.

Keywords: genetic tree improvement

nursery operations

tree phenology

growth

Abstract: Containerized Douglas fir (Pseudotsuga menziesii) seedlings from warm coastal and cool mountainous Oregon seed sources, grown under natural conditions, were chilled at constant temperatures of 5, 7 or 9 degrees C for 9, 11, 13 or 15 weeks beginning in mid-October. After a growth period of 9 weeks following chilling, degree of budbreak and dry weight of new shoot growth were measured. The longest and coldest chilling treatment produced the greatest growth response for all seed sources. Results are discussed with reference to predicted global warming, in particular the potential difficulty of reducing Douglas fir chilling requirements through tree breeding programmes.

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32. McKay, H.M. and A.D. Milner. 2000. Species and seasonal variability in the sensitivity of seedling conifer roots to drying and rough handling. Forestry-Oxford 73(3): 259-270.

Keywords: planting operations

tree physiology

tree/stand health

tree phenology

Abstract: The ability of the fine root system of 2-year-old bare-rooted planting stock of Picea sitchensis of Queen Charlotte Islands (British Columbia, Canada), Oregon and Alaskan (USA) provenances, Pseudotsuga menziesii, Larix kaempferi and Pinus sylvestris to withstand standard drying and rough handling treatments was tested at regular intervals from September 1990 until April 1991, at a nursery in NE England. Details are given of nursery treatments (sowing in spring 1989; undercutting in year 2 in June-mid-August, depending on species, and wrenching at 2-wk intervals until mid-October; and lifting at 2-wk intervals from September-November 1990 to April 1991). Electrolyte leakage was used to quantify the damage to the fine roots. Stressed seedlings had significantly greater leakage values than untreated seedlings and dried seedlings had significantly greater leakage values than roughly handled seedlings, but the responses varied with species and lifting date. After drying, leakage values increased in the order P. sitchensis (Oregon and Queen Charlotte Islands), P. sylvestris, L. kaempferi, P. sitchensis (Alaskan), P. menziesii. Leakage values after rough handling increased in the order P. sitchensis (Alaskan and Queen Charlotte islands), L. kaempferi, P. sitchensis (Oregon), P. sylvestris, P. menziesii. Leakage values were greatest in early September. During September and October there was a rapid downward trend in leakage. From December to February, leakage values decreased very slowly. In March leakage values of untreated and roughly handled seedlings increased slightly but decreased further in dried seedlings. These results indicate that great care must be taken during autumn planting to protect bare-rooted seedlings, particularly P. menziesii, from drying and to a lesser extent rough handling. Fine root leakage values following drying and rough handling had a significant linear relationship with the logarithm of the number of days required for the terminal bud to burst.

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