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Genetics of Douglas-Fir

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Genetics of Douglas-Fir Historic, archived document Do not assume content reflects current scientific knowledge, policies, or practices. \ of DOUGLAS-FIR • • • • • • • • •• • • • • • • • • • • U.S. Department of A . Forest Service grrculture This publication is one in a series on the genetics of important forest trees of North America being published by the Forest Serv­ ice, U.S. Department of Agriculture, in cooperation with the Society of American Foresters. Development of this series is in accord with the resolutions of the World Consultation on Forest Genetics and Tree Improvement at Stockholm in 1963 and the Second World Consultation on Forest Tree Breeding at Washington, D.C., in 1969. The Committee on Forest Tree Improvement of the Society of American Foresters undertook the preparation of manuscripts for North American species. CONTENTS Page RESEARCH SUMMARY -------------------------------­ 111 INTRODUCTION -----------------------------------__ _ 1 THE GENUS --------------------------------------- __ _ 2 Taxonomy ----------------------------------------­ 2 Cytology ------------------------------------------­ 2 Ancestral Distribution ----------------------------__ _ 2 THE SPECIES IN NATURE ___________________________ _ 3 Present Distribution ------------------------------ __ _ 3 Habitat -------------------------------------------­ 4 Growth -------------------------------------------­ 5 GENETICS -------------------------------------------- 5 Crossability ------------------------------------- __ _ 5 Genetic Markers -----------------------------------­ 5 Reproductive Development ___________________________ 6 Flowering ____________________________________ _ 8 Seed -----------------------------------------­ 9 Variation - -------------------------------- --------­ 10 Survival --------------------------------------­ 10 Growth ---------------------------------------­ 10 Form ------------------------------------------ 13 Phenology ----------------------------------- __ _ 15 Resistances -----------------------------------­ 15 Physiological Variation -------------------------- 16 VVood -----------------------------------------­ 17 Age of Trait Expression ____________________________ _ 17 GENETIC TECHNIQUES ------------------------------­ 18 Selection ------------------------------------------­ 18 Growth ---------------------------------------­ 18 Form ------------------------------------------ 19 Phenology _-----------------------------------­ 19 Resistances ---------------------------------·--­ 19 Cone Production -----------------------------------­ 19 Controlled Pollination ------------------------------­ 19 Testing -------------------------------------------­ 20 Asexual Reproduction ------------------------------­ 21 Grafting --------------------------------------­ 21 Rooting ---------------------------------------­ 22 APPLIED PROGRAMS ---------------------------------·­ 22 History -------------------------------------------­ 22 Seed Certification ----------------------------------­ 23 Seed Production Areas ------------------------------­ 23 Seed Orchards -------------------------------------­ 23 VVide Crossings ------------------------------------­ 24 Clonal Progra-ms -----------------------------------­ 24 Progressive Programs ------------------------------­ 24 STRATEGIES ----------------------------------------­ 26 Sources of Variation -------------------------------­ 26 Gains --------------------------------------------­ 26 Adaptation ----------------------------------------­ 27 ACKNOWLEDGMENTS -------------------------------­ 27 LITERATURE CITED ---------------------------------­ 28 i RESEARCH SUMMARY Douglas-fir (Pseudotsuga menziesii [Mirb.] Details of the life cycle of Douglas-fir have Franco) dominates the most productive forest only recently become complete enough that the lands of Western North America. More genetics germinal line can be followed through pollen research is being done on this species than on and seed development. Pollen is captured by a any of its associates. Tree improvement, how­ sea-anemone-like growth of the developing seed. ever, has lagged because of technical problems The species is characterized by very large, for which solutions were only recently found. round, featureless pollen grains with a me­ Evolution of the genus is relatively recent, chanically strong intine layer permitting expan­ possibly originating from Larix which it closely sion longitudinally to the 500 micrometers resembles in morphology and floral mecha­ needed to grow across a liquidless micropylar nisms. The species existed north of its present canal. No physiological or physical barriers to range in recent geological times. Present distri­ pollination are known, but embryo collapse fol­ bution of the six species of Pseudotsuga around lowing fertilization is common. One embryo the north Pacific Rim places four species in per seed is normally produced, the competitive Asia and two in North America. Only P. men­ outcome of up to nine pollen grains per micro­ ziesii has a chromosome complement of N = 13 pylar canal, four to six archegonia, and up to instead of N == 12. Only crosses between P. four cells that can contribute their chromo­ menziesii and P. macrocarpa have succeeded. somes to the resulting embryo. Seedlings with P. menziesii's latitudinal distribution from 19° chlorophyll-deficient recessive marker genes and to 55 ° N. is the most extensive of commercial yewlike mutations are reported. Polyploidy has western conifers. Most of its traits display clinal been induced with colchicine. variation along its range, which resembles an The species grafts well initially, but about inverted V. A variety, glauca, is recognized as 35-percent rejection of grafts is normal. Early applying to more continental races of the in­ recognition of anatomical symptoms permits terior arm. Maximum development occurs along two kinds of evasion techniques so seed or­ its coastal arm west of the Cascade Range and chards may now be established virtually free the Coast Ranges. The species occurs on almost of incompatible grafts. A high percentage of any moist, well-drained forest habitat in its cuttings from year-old seedlings root, but suc­ range below midalpine zones, yet it withstands cess falls off to about 20 percent for 10- to droughts of several months. Its evolutionary 50-year-old trees. Cuttings of mature trees are niche as a fire species arises from its rapid usually rooted with great difficulty. growth, tallness, durable wood, thick bark, and Genetic variation is documented for many long life, the last two traits being of minor traits. Exact adaptation to local environments breeding interest. Its rapid growth period is is suggested from many studies showing growth, preceded by a decade of slow seedling growth. phenological, and terpene differences between The older growth, soft, fine-grained wood pro­ nearby populations despite pollen exchange. In­ duction, is preceded by a long period of pro­ herent growth differences appear to follow a ducing coarse-grained wood. westwide pattern of relative assurance of ade­ Douglas-fir is monoecious. It rarely flowers quate springtime moisture. The best inherent as early as 2 years and produces only small growth has evolved in the western and northern quantities of seed the first decade. Cone crops parts of Douglas-fir's range and the poorest in are usually cyclic, causing occasional severe the eastern and southern parts. Growth ratios local seed shortages. Nitrate fertilizers and gib­ between families within local populations can berellins have been successfully used to enhance exceed 2 :1, providing impetus for practical im­ crops. Naturally harvested cones average about provement of already adapted genotypes. 16 filled seed but can range up to 54 filled seed Growth traits require one or more decades for on some trees. Only about 7-percent average adequate expression. Heritabilities of growth selfing occurs despite the majority of pollen and form have been generally moderate and being received from the tree itself. Inbreeding heritabilities of phenology, resistances, and coefficient is estimated at 0.025, leading to wood traits generally strong. estimates of 1 to 1.5 percent inbreeding depres­ Practical programs include seed orchards, sion in seedling height. wide crossings, wind-pollinated seed, and clonal iii emphasis. Practically all programs in the Doug­ generation programs for full-sib crosses are las-fir region use locally adapted parentage. established at several seedling orchards. Varia­ Programs in other temperate forest regions tions of the commonly used technologies have involve search for adapted races. Earlier seed developed especially for Douglas-fir. Special orchard problems are now largely alleviated. A strategies have also been developed to maxi­ graftless concept, based on tests of large num­ mize efficiencies, to utilize within-stand or racial bers of parents with wind-pollinated seed, is variation, and to minimize losses from mala­ widely used in the Douglas-fir region. Second- daptation. iv GENETICS OF DOUGLAS-FIR Roy R. Silen 1 INTRODUCTION In the natural forest, Douglas-fir (Pseudot­ Pinaceae is interesting. The problems of graft­ suga menziesii [Mirb.] Franco) is a remark­ ing incompatibility, pollen contamination, dif­ ably successful species. It vies with coastal ficult rooting, and inadequate cone production redwood for height supremacy among the of Douglas-fir delayed large scale improvement world's conifer s. In the droughty, fire-prone programs until recent research found ways to
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