w < 72. . 3:1, TH '1" W 31“ HIGH Ill/g ”LI/'BTARIES lllllllllllll This is to certify that the thesis entitled Evaluation on Height and Diameter of 9-year-old Progeny Test of Native Aspens and Their Hybrids in Michigan presented by Putranto Budiono-Agung Nugroho has been accepted towards fulfillment of the requirements for Master Forestry degree in Major professor Date F649 //r1 [990? 0-7 639 MS U is an Affirmative Action/Equal Opportunity Institution LIBRARY i Michigan State University PLACE IN RETURN BOX to remove this checkout from your record. TO AVOID FINES return on or before date due. DATE DU SW] Li L_J i:i__i- i—l :ji—i l:L_J Ei—l: —T[_ l— i— l l MSU Ie An Affirmative Action/Equal Opportunity Institution oWMS-M EVALUATION ON HEIGHT AND DIAMETER OF 9-YEAR-OLD PROGENY TEST OF NATIVE ASPENS AND THEIR HYBRIDS IN MICHIGAN BY Putranto Budiono-Agung Nugroho A THESIS Submitted to Michigan State University in partial fulfillment of the requirement for the degree of MASTER OF SCIENCE Department of Forestry 1992 /'/;" ~‘ f7f— //V:,J\ ABSTRACT EVALUATION ON HEIGHT AND DIAMETER OF 9-YEAR-OLD PROGENY TEST OF NATIVE ASPENS AND THEIR HYBRIDS IN MICHIGAN BY Putranto Budiono-Agung Nugroho An evaluation on trembling aspen (Populus trgmulgides), bigtooth aspen (Egpglgg gzangigentata) and their hybrids was undertaken at Water Quality Research Center, Ingham co., Michigan. Progenies of open- and controlled-pollination were generated from trees from Upper and Lower Peninsula. The progenies were planted in two-tree plot in a randomized complete block design with six replications. The F1 hybrid of trembling and bigtooth aspen was able to establish as fast as their parents. Height and stem diameter of the hybrids was intermediate at the first- and second-year but superior at 7 and 9 years. Progenies with female parent from Lower Peninsula, especially central Lower Peninsula, grew faster than those from Upper Peninsula. Height and diameter was highly correlated. Age-age correla- tion among ages 1, 2, 7 and 9 years were significant. An analysis using North Carolina I mating design identified a relatively high genetic variation. The compo- sition of additive and dominance variance changed with age. At 7 and 9 years the genetic variance was dominated by the dominance variance. To God, The Almighty Thank you. ACKNO'LEDGMENT I wish to express my sincere gratitude to my major professor, Dr. J.W.hanover, for his guidance and patience during the course of my graduate education. I would also like to express my appreciation to my graduate committee, Dr. D.E. Keathley and Dr. R. Ward, and to Dr. D. I. Dickmann for their helpful suggestions. I owe a debt of appreciation to all MICHCOTIP staff and graduate students who generously gave their time to help me during the completion of my study. A special thank goes to my parents, Kridarso Budi- Prajitno and Hajat Radliani, and to my fiancee, Era Retnoningsih, for their unending support and encouragement throughout my study. Finally, I am indebted to all Indonesian people for their sacrifices in providing financial support which were instrumental in enabling me to pursue a personal goal. TABLE OF CONTENTS page LIST OF TABLES ... ..... .................................v LIST OF FIGURES . ........ .. ...... ......................vii INTRODUCTION ........... ........... .....................1 LITERATURE REVIEW ................................ ..... 5 - Aspen Distribution and Variation ................5 - Aspen Hybrids ....................................9 - Age-age correlation eeeeeeeeeoeeeeeeeeeeeoooeee014 MATERIALANDWTHODS ......OOOOOOO0.00.00.00.0000000000016 Progeny Production ..............................16 Plantation Establishment .......................18 Data Collection ................................19 - Data Analysis ..................................19 l. Phenotypic performance among taxa .........l9 2. Geographic significance of parent .........21 3. Heritability and variance component estimates .................................22 4. Correlation ..............................26 RESULTS AND DISCUSSION ......OOOOOOOOOOOOOOOOOO00......27 - Phenotypic Performance among Taxa ...............27 1. Survival rate ............................27 2. Stem diameter and height...................27 - Geographic Significance of Parent ...............4O - Heritability and Variance Component Estimate ....47 - Correlations ....................................57 1. Stem diameter - height correlation .......57 2. Age - age correlation ....................60 CONCLUSION ............................................64 RECOMENDATIONS .........................................67 LIST OF REFFERENCES ....................................7l APPENDICES iv Table Table Table Table Table Table Table Table Table Table 9. LIST OF TABLES page Form of analysis of variance (ANOVA) for analyzing height and stem diameter growth of taxa and families within taxa, based on tree-plot unit data ............................20 Form of analysis of variance (ANOVA) for analyzing effect of geographic of parental origin OIO00.0.0.0.........OOOOOOOCOOOOOOOO00.0.22 Form of ANOVA and EMS of NC mating design I for analyzing male and female parent and variance component estimate of all population (base on plot mean data) .......................24 Form of ANOVA and its expected mean square for analyzing within plot component of variance (based on individual-tree data) ................25 Formula for calculating narrow-sense heritabilities, utilizing variance components estimates derived from NC I mating design......25 Survival rate of each aspen taxon at ages 1, 2, 7 and 9 years (in percentages) .........27 F value of analysis of variance of stem diameter and height among taxa and among family within taxa ...O0....0......0..........OOOOOOOOOOOOOOOOZB Means of stem diameter of each aspen taxon at ages 1, 2, 7 and 9 years.....................31 Means of height of each aspen taxon at ages 1’ 2' 7’ andgyears CO...00.0.0000000000000000032 10. Number of families of each aspen taxon that. comprises 15 (10%) best families in height and stem diameter at ages 1, 2, 7 and 9 years ....36 Table Table Table Table Table Table Table Table Table Table Table Table Table Table 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. A1. A2. vi page Stem diameter (cm) and percentage ranking of the best family and the best individual of each aspen taxon at ages 1, 2, 7 and 9 years ...................................38 Height (cm) and pencentage ranking of the best family and the best individual of each aspen taxon at ages 1, 2, and 7 years ...................................39 F value of analysis of variance of height and stem diameter among maternal parent origin.(Entries mare cans of families at each region) ...............................41 Mean of stem diameter of aspen families among geographic areas at ages 1, 2, 7 and 9 years...43 Mean of height of aspen families among geographic area at ages 1, 2 and 7 years .....43 The fifteen best aspen families (in diameter) at ageg year.C............0.0.0.000000000000045 The fifteen best aspen families (in height) at age?year.........OOOOOOOOOOOO00.000.000.0000046 F value of ANOVA derived from the nested design (NC design 1) at Table 3. ..............48 Components of variance of stem diameter and height derived from NC 1 mating design......49 Genetic and environmental variance and narrow-sense heritability of stem diameter and height derived from NC 1 mating design ...52 Coefficient of correlation between stem diameter and height at ages 1, 2, and 7 years 00............OOOOOOOOOOOOOOOOO00.0.0058 Phenotypic coefficient of correlation of stem diameter and height among ages 1, 2, 73nd9years O.......OOOOOOOOOOOOOOOOOO ....... 62 Families that at age 9 years at least represent at four replications Families that were used for NC-l mating design Figure Figure Figure Figure Figure Figure Figure Figure Figure 1. 2. 3. 4. 5. 6. LIST OF FIGURES page The location of seed and catkin-bearing branch collections........................ ..... 17 Means of stem diameter at ages 1, 2, 7 and gyears ........ ......OOOOOOOOOOOOOOOOOO ....... 33 Means of height at ages 1, 2 and 7 years.......33 Means of stem diameter among geograpic areas at ages 1, 2, 7 and 9 years ...................44 Means of height among geographic areas at89381' zand7years .00000000000000000000044 Components of variance of stem diameter at ages 1, 2, 7 and 9 years derived from NC I mating design (in percentage).............So Components of variance of height at ages 1, 2, and 7 years derived from NC I mating design (in percentage) ........................50 Genetic and environmental variance estimates of stem diameter at ages 1, 2, 7 and 9 years (in percentage)... ..... ..................53 Genetic and environmental variance estimates of height at ages 1, 2 and 7 years (in percentage)......OOOOOOOOOOOOO0.0.0.000000053 Figure 10. Family- and individual tree-based heritability of stem diameter at ages 1, 2, 7 andgyears ......OOOOOOOOOOOO00.0.0000000054 Figure 11. Family- and individual tree-based heritability of height at ages 1, 2 and 7years ......OOOOOIOOOOOOOOO......OOOOOOOOOOOS4 Figure 12. Coefficient of correlation between stem diameter and height at ages 1, 2 and 7 years..59 vii viii page Figure 13. Age-age coefficient of correlation of stem diameter and height among ages 1, 2, 7 and 9years COC0.0.0.0.0....OOOOOOOOOOOOOO00.0.000063 13189229129! Trembling asnen (Penning treaulgideei and bigtooth aspen (Renulus grandidentata) are dominant species throughout the western Great Lakes area (Henry 8 Barnes 1977). These species grow sympatrically in Michigan and over a large portion of northeastern United State adjacent to Canada (Pregitzer & Barnes 1980). Aspens are recognized as multiple-use species. They produce forage and cover for domestic livestock and wildlife, produce wood fiber for pulp, form protective firebreaks, yield high-quality of water, and are aesthetically attractive in the landscape (Debyle and Winokur 19..). Aspen is the most important pulpwood species in Michigan (Blyth & Smith 1982). However, the use of aspen in plantations has been avoided because genetically improved planting stock is not readily available (Reighard 1984). Another reason is the difficulty of cloning superior. genotype or producing adequate seed for commercial nurseries (Dickmann 1992, personal communication). Genetic variation of aspen is quite large. Large. variation exists in height/diameter.growth rate (Zahner & Crawford 1965, Barnes 1969, Einsphar & Benson 1967, Mitton 8 Grant 1980), survival rate (Pauley 1963, Pauley et a1. 1963) photoperiod responce (Vaartaja 1960), phenology (Barnes 1969, Brissette 8 Barnes 1984), specific gravity, fiber length, and chemical content of wood (Buijtenen et a1. 1959, Einsphar 8 Benson 1967), suckering ability of roots (Schier 8 Chambell 1980, Barnes 1969), rooting ability (Schier 1974, Schier 8 Chambell 1980) and susceptibility to diseases and insects (Wall 1971, Copony 8 ' Barnes 1974). Johnson (1942) had reported that selection for height and diameter growth appeared to have lack effect on wood quality. This lack of correlation might increase the efficiency of selection for these traits. Trembling aspen and bigtooth aspen are easy to hybridize (Henry 8 Barnes 1977). Natural hybridization of these species has been reported by many authors (McComb 8 Hanson 1954, Einsphar 8 Joranson 1960, Andrejak 8 Barnes 1969, Wagner 1970). Natural hybrids were commonly found in extensively disturbed areas and grew together with their parents (Andrejak 8 Barnes 1969, Wagner 1970, Henry 8 Barnes 1977). Artificial hybrids were first produced by Heimburger (1936) and later by Pauley (1963). The ease of hybridization might allow the possibility of enriching variation of these species with new sources of germplasm. The new germplasm may or may not be desirable. It is 3 desired if it has beneficial effects, such as faster growth or more resistance to disease, but undesired if give the opposite effects. Comparison between native aspen and their hybrids has been reported in flowering phenology (Pregitzer 8 Barnes, 1980), germination rate (Henry 8 Barnes 1977), and initial height 8 diameter growth (Henry 8 Barnes 1977, Brissette 8 Barnes 1984, Einsphar 8 Benson 1964, Reighard 1984). Various results from different studies of initial growth have been reported. Henry 8 Barnes (1977) reported that during the first 4 months hybrid seedlings grew faster than the progeny of either their parent, but at 6 months-old B.§zgmglgidgs was leading, followed by the hybrids, then £.g:andidgntata. Brissette 8 Barnes (1984) found that at the end of the first year B.§zgmglgidg§ was the tallest followed by the hybrids then 2.9:and1dgntata, but at the end of the second year 2.g;and1dgntg§a was second and the hybrid was the shorthest. A study by Reighard (1984) showed that at the first and second year measurements the hybrid was leading the progeny of either parent. Another study by Einsphar 8 Benson (1964) showed that, at one and four years old, £.§:gmglgidg§ was the tallest followed by the hybrids, then £.grand1dgn§atg. The increasing demands for aspens by the forest products industry have developed because of new technology to use aspen for products other than the traditional pulp 4 and paper (Reighard 1984). The potency of aspen for genetic improvement (ease of hybridization and large genetic variation) and the lack of genetically improved planting stock for plantations are challenges for breeders to initiate an improvement program for this species. In 1982 Michigan State University established a progeny test of aspen and their hybrids in Michigan. Many years will be required for testing and screening superior genotypes of these species to be released to the public as improved stock. However, by developing an early evaluation, improved stock could be ready for the release in the near future. This study reports the 9-year-old results of a progeny test plantation in the MSU Water Quality Research Area, Ingham Co. Michigan. The objectives of this study are: 1. To identify the fast-growing (height and stem diameter) aspen families and their hybrids. 2. To quantify genetic variation in the native aspen families and their hybrids. 3. To analyze age-age correlations of growth (height and stem diameter) of aspen families and their hybrids. TU V o V o . 119911195 treauleides and 292212: W are the only two of four native poplar in Michigan referred to as aspen (Graham et al. 1963). Trembling aspen (2922135 txemglgidgg) is a boreal-temperate species and bigtooth aspen (£92313: grandidentata) is a temperate mesic species (Fowell 1965). Trembling aspen is widely distributed on the North American continent. It grows from Alaska to northern Mexico, but is found mostly in north mid-western United States (Dickmann and Stuart, 1983). The southern range of its distribution extends along the Appalachian mountains to Georgia (Strotham and Zasada 1957). This species is more adaptable than bigtooth aspen (Graham et al.1963). It grows on more varieties of soil, but its growth is most satisfactory in well-drained loamy soils, and in land with a water table within 1.5 m of the surface (Dickmann and Stuart 1983, Graham et al. 1963). The distribution of bigtooth aspen is much more restricted. It ranges from Maine and southern Canada to Tennessee and North Carolina (Graham et a1. 1963). Bigtooth aspen is most often found in well-drained medium 6 to coarse texture upland soils (Dickmann and Stuart 1983, Graham et al. 1963). When bigtooth and trembling aspen are growing together, bigtooth aspen often outgrows trembling aspen, however, due to its greater susceptibility to some juvenile diseases, bigtooth aspen is not likely to become established in some places (Graham et a1. 1963). Since trembling aspen and bigtooth aspen grow in a relatively wide range of environments, the existence of genetic variation can be expected. Some studies on natural variation have been reported from various authors. Pauley (1963), studying aspen seedlings from various seed sources in Massachusetts, reported evidence of geographic variation on survival rate and growth. Survival rate and growth of seedlings from Lake states origin was similar to those from New England, but western seedlings from Arizona to the Yukon territory were weak and almost died by the age of 12 years. Brissete 8 Barnes (1984), comparing phenology and growth of trembling aspen from Michigan and western North Americans growing in southeastern Michigan, documented that seedlings from western North American origin stopped growing earlier than those from Michigan. He also reported that after 2 years, the average height of the western progeny was only 26-38 percent of the height of Michigan progenies. The poor performance of western seedling origin was probably due to the problem of adaptation to 7 photoperiod or temperature (Spurr 8 Barnes 1980, Brissete and Barnes 1984, Reighard 1984). Vartaaja (1960) found that daylength of the different latitudes was important for growth. Comparing seedlings from Wisconsin and northern Saskatchewan, he found that growth response to short-day condition between those sources of seedlings was very different. Brissete and Barnes (1984) reported that, in a A high daily mean temperature condition, aspen from lower temperature showed low rate of photosynthesis and high rate of respiration. Einsphar and Benson (1967), studying geographic variation of trembling aspen in Wisconsin and the Upper Peninsula of Michigan, reported a well-defined south-to- north trend of decreasing specific gravity of wood. Highly significant differences for a number of important growth and wood properties were obtained between clones within stand and stand within areas. A similar result was also reported by Barnes (1975) from a study of phenotype of leaves representing western aspen from southern Utah and Colorado northward to the Canadian border. A tendency of smaller and narrower leaves following a clinal pattern from south to north was evident. Einsphar, et al. (1963), studying natural variation in triploid aspen, reported that differences in tree volume, specific gravity, fiber length, fiber strength, crown volume, leaf size and shape appeared to be controlled 8 genetically. A great deal of variation within areas was found, but differences between areas were more striking. Several studies have also been carried out to document clonal variation in aspen. From various studies, Debyle 8 Winokur (19..) has compiled evidences of clonal variation of aspen for several characteristics, such as annual height and diameter growth, bursting of floral buds, timing of leaf flushing, suckering capacity, rooting, susceptibility to diseases and insects, flowering time and branching habit. Barnes (1969), for example, identifying clones from populations in the northern Lower Peninsula of Michigan, reported that aspen exhibits an enormous amount of inter clonal variation, even in local populations on fragmentary parts of their range. Noticeable differences between clones occurred in growth rate, clone profile, density of ramets, and suckering ability. He also documented extensive inter- clonal variation in leaf morphology, size and shape. However, in some instances, intra-clonal leaf variation was more striking and greater than inter-clonal variation. Mitton and Grant (1980), in the Colorado Front Range, found a significant positive correlation between clone heterozygosity and annual diameter growth. Another study by Zahner and Crawford (1965) documented large differences in growth of adjacent clones on the same site. 9 Genetic variation in other characteristics has also been studied. For example, Wall et al. (1971) noted that, in Manitoba, some clones became chlorotic on nutrient- deficient sites while other clones did not. Copony and Barnes (1974), studying four different sites in Michigan, reported that susceptibility to flypgxylgn canker varied markedly among clones. Tew (1970) found different carbohydrate reserves in roots between aspen clones. AW. Trembling aspen crosses readily with other species from the genus Populus within section Leuce, producing viable hybrids (DeByle and Winokur 19..). Successful crossing involving trembling aspen or bigtooth aspen as a parent, such as crossing with Populus alga, B.§1g§glg11, Redeem. B-triszhszsarna. Humanism: and Fireman. has been reported (Zsuffa 1973, Dickmann and Stuart 1983). Natural hybrids between trembling aspen and bigtooth aspen was first reported and described by Victorin (Barnes 1961). Although the flowering time of trembling aspen was earlier than that of bigtooth aspen, natural hybrids of trembling aspen and bigtooth aspen were not infrequent (Heimburger 1940, Pauley 1956). In central and eastern Massachusetts, Pauley (1956) observed scattered hybrid individuals and hybrid swarms of these species. In Michigan, natural hybrids of these species are not 10 uncommon. Between 1956 and 1960, Barnes (1961) discovered natural hybrids of trembling and bigtooth aspen in 10 counties of Michigan. He found that the hybrid was apparently mush more abundant in south-eastern Michigan than in the northern tip of the Lower Peninsula. This phenomenon might be caused by a condition, in an area of temperature inversion, where the flowering of female trembling aspen was retarded until it corresponded with the flowering time of neighboring bigtooth aspen (Pauley 1956). Trembling aspen and bigtooth aspen are also easy to hybridize artificially. Einsphar and Benson (1964) described a simple procedure of hybridization technique known as the "cut-branch technique." The procedure is as follows: first, flower buds from the trees to be crossed were collected. The male flower buds are forced by placing the branch collection in a vase of tap water. The water is changed daily and a small disk from the end of each branch is clipped. After seven to ten days (at 65°F) the pollen will be already available to be collected and is then stored over calcium chloride at 38° to 40°F. Female branch collections are handled in a similar manner with the exception that after pollination, which is accomplished by applying the pollen with a small brush, the collections are placed in ice water to reduce the possibility of bacterial plugging. Then in 21 to 24 days the seed can collected and separated from the attached cotton. 11 Many experiments involving trembling aspen and bigtooth aspen hybrid had been reported (Heimburger 1940, Pauley et al. 1963a, Einspahr 8 Benson 1964, Henry 8 Barnes 1977, Brissete 8 Barnes 1984, Reighard 1984). Unfortunately only early growth analysis was available in most cases. .The earliest experiment involving aspen hybrid conducted in the United States was initiated in 1924 by Oxford Paper Company of Maine, under direction of A. B. Stout and E. J Schreiner (Einsphar 8 Benson 1964, Dickmann and Stuart 1983). The experiment included parental trees of three white poplars, five aspens, nine balsam poplars and seventeen black poplars and cottonwood (Dickmann and Stuart 1983). Unfortunately, there was no further report on the aspen of this experiment. In 1935, Heimburger (1940), at Petawawa Forest Experiment Station of Ontario Canada, successfully crossed trembling aspen and bigtooth aspen. The objective of this experiment was to produce hybrid aspen suitable for reforestation in Ontario. The breeding goal included fast growth, hardiness, resistance to insects and disease, and improving rooting ability of stem cuttings (Dickmann and Stuart 1983). In this experiment Heimburger also crossed trembling aspen and bigtooth aspen with gray poplar (2.x gangsggns Sm). In 1937 the experiment showed that trembling aspen x bigtooth aspen hybrids had a good survival rate and a fair degree of resistance to rust 12 caused by uglgmpggrg sp, while bigtooth aspen x gray poplar hybrids had already perished (Heimburger 1940). In 1954, an industry-sponsored program for the improvement of aspens in Lake states was initiated at the Institute Paper Chemistry of Appleton, Wisconsin, under direction of P. Joranson and D. Einsphar (Dickmann and Stuart 1983, Einsphar 8 Benson 1964). This program included selection of plus trees from natural stands, hybridization, and created polyploidy aspen to produce vigorous growth and better wood quality (Dickmann and Stuart 1983). At the early growth stage, much variation between crosses appeared in growth, tree form, and wood properties (Einsphar and Benson, 1964). The early growth of bigtooth aspen was relatively slower than either trembling aspen x bigtooth aspen hybrids or triploid hybrid produced by crossing natural diploid trembling aspen with tetraploid european aspen (£.t:gmgla). The triploid hybrid grew more vigorously than the diploid trembling aspen. From the same experiment, Benson 8 Einsphar (1967), comparing 4 years growth of triploid trembling aspen clones, triploid trembling aspen x european aspen hybrid, and diploid trembling aspen, reported a significant difference among taxa on some traits, such as on growth, specific gravity of wood, natural pruning, number of branches, branch angle, stem straightness and branch length. 13 Henry and Barnes (1977), comparing reproductive ability of bigtooth aspen, trembling aspen, and their hybrids in southern Michigan, reported that in some reproductive traits the hybrids were at lower degrees than the progeny of either their parent but intermediate in others. Seed production per shoot and seed germination of the hybrids were significantly lower than those of their parents. Initial height growth of the hybrid at four weeks was greater than the progeny of their parents. At eighteen weeks the hybrid is significantly lower than the progeny of trembling aspen but not significantly different from the progeny of bigtooth aspen. Following height growth studied by Henry 8 Barnes (1977), Brissete and Barnes (1984) recorded that, at one year, trembling aspen were significantly taller than bigtooth aspen and their hybrid. The hybrid was in the middle and bigtooth aspen was the lowest. At second year, trembling aspen was still significantly taller than the others, but there was no significant difference between the hybrid and bigtooth aspen. This result might indicate that, at early growth, hybrids of bigtooth aspen and trembling aspen showed neither hybrid vigor nor marked growth inferiority, compared to their parents. Brissete and Barnes (1984) also investigated inter- and intra-specific hybridization between trembling aspen from Utah and trembling aspen and bigtooth aspen native to 14 Michigan. They found that, at second year, the progeny of bigtooth aspen from Michigan crossed to trembling aspen from Utah was only 64% as tall as those crosses between Michigan aspens. Crosses between trembling aspen from Utah and those from Michigan were only 83% of mean height of those from Michigan. This indicates the importance of parental sources for hybridization for a specific location. Reighard (1984), studying a two-year-old progeny test of trembling aspen, bigtooth aspen and their hybrids at five location in both peninsulas of Michigan, reported that maternal parent affected autumn leaf color, branchiness and bud morphology, more than paternal parent. The phenology of the hybrids, compared to their parents, was intermediate, but they suffered from a number of disorders more than their parents. The test also showed a significant difference in height, basal diameter and biomass production. Growth of trembling aspen was above bigtooth aspen and their hybrids. The growth increased with the latitude of plantation site. However, most of the hybrid's families had growth rate below the plantation means. e- c a o . The goal of an advance-generation tree breeding program is to maximize gain achieved per unit time (Zobel and Talbert, 1984). The breeder can increase efficiency of the overall program by eliminating poor trees and 15 concentrating on potential superior trees. Therefore, early selection must be considered. If early selection is to be successful, there must be evidence of high correlation between performance at rotation age and at a younger age. Unfortunately, juvenile- mature correlation in forest trees is usually low (Zobel and Talbert, 1984). The low juvenile-mature correlation has been reported in loblolly pines, ponderosa pines, Douglas fir, western white pines, slash pines, black walnut and various hardwood species (Wright 1976). However, Mohrdiek (1979), evaluating 15 F3 Leuce progeny trial among crosses of 299913; tzgmulg, 2. alga, Box camps Sm. BMW and B-mndidenteta from age one year to age twenty years, had reported high correlation of phenotypic growth across ages, he found that the age-age coefficient of correlation between age 20 years and ages 15, 11, 9, 3, 2, 1 years were 0.952, 0.934, 0.828, 0.554, 0.483 and 0.462, respectively. Mohrdiek also suggested that a test interval of eight years seems to be sufficient for’Fg Leuce progeny. Reighard (1984), measuring age-age coefficient of correlation between nursery height growth at one year and height growth after two growing season in the plantation, reported r s 0.48. He indicated that, with this correlation coefficient, an early selection of’F} progeny of aspen would be possible. groggny Productigg Progenies were produced by G. Reighard and the Michigan Cooperative Tree Improvement Program (MICHCOTIP) at MSU in 1981. Seeds and catkin-bearing branches of bigtooth and trembling aspen were collected during March and April 1979 and 1980 from most counties in both . peninsulas of Michigan (Figure. 1). Similar material of a putative white poplar-bigtooth aspen hybrid (2.x Igulgauiana) from southern lower peninsula were also collected. Controlled pollination was done using cut—branch technique (Einsphar 8 Benson 1964). In addition to the seed obtained from open pollination, the progeny of controlled pollination represented 48 half-sib and 66 full-sib families of both bigtooth and trembling aspen, 72 full sib families of hybrid asraen (2.x smithii -= Lgrandidentata x 2.;rgmglgidgg, reciprocally), and 20 full sib families of crosses of bigtooth and trembling aspen to the putative white poplar-bigtooth aspen hybrid. Seed was sown in the nursery on May 26 1981. Cultural procedures similar to those mentioned in "Aspen seedling production in a commercial nursery" (Benson 8 Dubey, 1972) 16 17 Figure 1. The location of seed and catkin-bearing branch collections 18 were used to grow the seedlings. Orthene was used to control insects. Seedlings were lifted the following March and placed in cold storage until the planting day. WM. Plantations were established on five sites, three in the Lower Peninsula and two in the Upper Peninsula. ' However, this study only deals with one plantation at the MSU Water Quality Research Center (Lower Peninsula, Ingham Co. Lat. 42.7N, Long 84.5W). The soil is well-drained and the texture is a fine sandy loam. The dominant vegetation on the site was grasses and perennial weeds. Planting was done in April and May 1982. Seedlings were planted by machine in two-tree plots with spacing 1.8 meter between trees within rows and 2.4 meter between rows. The experimental design was a randomized complete block with six replications. Site preparation consisted of mowing the existing vegetation with rotary mowed in August 1981 and spraying seven liters/ha of glyphosate in one-meter-wide strips three to four weeks later. At the following spring planting, 2.8 kg/ha of Simazine was applied over the tops of the seedlings and on to the glyphosate-sprayed strips. To control the invading grasses, Glyphosate was spot- sprayed once in 1982. Mowing was done once during each of the following years. 19 mm- Data were collected for survival rate, stem diameter and height. Stem diameter was measured in 1982 and 1983 (at five cm above ground), and in 1988 and 1990 (at breast height) to the nearest 2.5 cm. Height was measured in 1982 and 1983 (to the nearest 5 cm) and 1988 (to the nearest 1 cm). Survival rate was recorded as percentage from the original number of trees planted. mum:- All analyses were done by using SAS Programs. Data were analyzed separately according to ages at measuring times. Since some trees were missing, all analyses, unless otherwise stated, utilize data from families that are at age 9 represented in least 4 replications. Analysis of variances and correlations were calculated for stem diameter and height at all ages measured. Survival rates of each taxon were simply presented as percentages calculated from number of survived trees at measuring times divided by number of trees initially planted. Height and stem diameter growth were subjected to an analysis of variance (ANOVA), based on the tree-plot units as entries, following a linear model as: 20 Yijk '-' u + R‘ + T] + Fun + Eiik where: YE“ = performance of plot-unit of kth family nested to jth taxa in the ith replication: u 2 overall mean; It a effect of ith replication: T - effect of j‘" taxa: i F - effect of the k‘" nested to j‘" taxa; kCl) E”k <= experimental error. Form of the analysis of variance (ANOVA) is presented in Table 1. Taxa were considered as fixed, while families within taxa and replications as random. Means of taxa were compared to each other by using Duncan's multiple range test. Table 1. Form of analysis of variance (ANOVA) model for analyzing height and stem diameter growth of taxa and families within taxa, based on tree-plot unit data. Sources de MS Fteet EMSV” Replicate. r-l MSR -- Taxa t-l us'r ns'r/nsu'r) v'. + rV'm, + er', Family t(f-l) nsrir) usr(r)/nss v5 + rvu“, within taxa Error (tf-l) MSE -- V3 (2'1) Total trf-l --- V r, t and f refer to number of replications, taxa and harmonic mean number of families/taxa respectively. 9 V'., V',,., and fV'. refer to variance error, variance family within taxa and variance among taxa, respectively. 21 2- Wr Effects of parental origin were analyzed for height and stem diameter. Geographic parental origins were simply grouped into four regions: 41.8 N to 43.0 N, 43.0 N to 44.2 N, 44.2 N to 45.4 N, and 45.4 N to 46.8 N. The first three regions represent parental trees from the Lower Peninsula, while the last one represents parental trees from the Upper Peninsula. Since the parents of the trihybrid were only from two regions, this taxa was excluded from the analysis. Analysis of variance was conducted for height and stem diameter following a linear model as: YUk = u + R§«+ Tj-+ zk-+IEUk where: Y," = performance of entries of jth taxa at kth regions in the ith replication: u = overall mean: R1 = effect of i‘" replication: Tj - effect of j‘" taxa: zk = effect of the kth regions. Eur = experimental error. Form of the ANOVA model is presented in Table 2. Both taxa and regions were considered to be fixed. Since numbers of trees of the treatments at each replication were different, analysis was conducted based on means of trees on the treatments (taxa and region) at each replication. 22 Contrast comparison tests were used for comparing trees with Upper Peninsulas parental origin to those from Lower Peninsula, while Duncan's multiple range test was used for comparing means of height and stem diameter trees among regions Table 2. Form of analysis of variance (ANOVA) for analyzing effect of geographic of parental origin. ‘ Sources Replicate :Taxa ; Regions , -UppervsLower j Peninsulas” : Error V Upper and Lower Peninsulas were compared using contrast comparison test. V t, z and r refer to number of taxa, regions and replications respectively. 3. v as c n n s e 9,- Since the population in this test is a mix of families (half-sib and full-sib families from controlled pollination, and half-sib families from open pollination), direct estimates of variance components and heritabilities cannot be done. Variance component and heritability were only approached from the controlled pollinated families together (regardless of taxa), by constructing a nested design (North Carolina design I). 213 After adjusting for missing families/trees, there were available 78 progenies of 27 male families that had been mated with at least 2 female families. Samples of 15 male families and 30 female families (in which each male was crossed into 2 females) were selected randomly without repetition. The progenies represent 30 families, 5 families of 2. grandidgntata, 9 families of 2. x smithii ' (2. grendidentefa x B. tremuleides). 6 families 2.x smithii (2. tggmulgidgg x B. grandidgntatg) and 10 families of 13- fungicides- An analysis of variance, by using the mating design, was conducted according to Becker (1984) following a linear model as: ij == u + Ri-itg + F *‘Ehn k(i) where: Y“, =8 plot-unit mean within the jth male parent and the k“ female parent: = overall mean: :- effect of 1th replication; do = effect of j“ male parent: re gs w s In» = effect of the kth female parent mated to jth male parent: E.. = environmental and remainder of genetic variance among plots. Replicate is considered as fixed while male parents and females within male parent as random. 24 Since some trees within plot were missing, an estimate of the individual variance within plots was obtained from individual trees by using the formula (Becker, 1984): Y“ =u+Pf+Efk where: Y“ = individual observation on kth individual of f“ plot: u = overall mean: Pf = effect of the f‘" plot; E“ - environmental and genetic variance among trees within plot The form of the ANOVA model and the expected means square (EMS) are presented at Table 3 8 4. Table 3. Form of ANOVA and EMS of NC mating design I for analyzing male and female parent and variance component estimate of all population (base on plot mean data) I Sources de Ms Ftest EMSVV Replicate r-l MSR -- Male m-l MSM MSM/MSF (M) V'.+rV'r(n)+er'. Female (male) m(f-1) MSF(M) MSF(M)/MSE Vfi+rVfl,, Male-female (mf-l) MSE -- V2 crosses x rep. (r-l) (error pooled) Total rmf-l MST V r, m and f refer to number of replications, male parent and number of female parent mating to male parent respectively. V V3, Vflm,and fVfi refer to variance error pooled, variance among female within male and variance among male parent, respectively 25 Table 4. Form of ANOVA and its expected mean square for analyzing within plot component of variance (based on individual-tree data) Ii Sources de SS MS EMsy Between rmf-l SS -- -— plot Within plot rfm(t-l) ss. usw v3 V r, m, f and t refer to number of replications, male parent and number of female parent mating to male parent and harmonic mean of number of trees per plot, respectively. V v1 variance among trees within plot. With the assumption of no occurrence of epistasis, the variance among male parent (Vfl) was considered equal to 1/4 of additive genetic variance, and variance among female within male parent (Vztm) was considered equal to 1/4 additive variance plus 1/4 dominance variance (Hallauer 8 Miranda 1981, Namkoong 1979). Then the narrow-sense heritability estimate was formulated according to Hallauer 8 Miranda (1981) as presented at Table 5. Table 5. Formula for calculating narrow-sense heritabilities utilizing variance components derived from NC I mating design. H H il Family based heritability Single tree based heritability 4 Vt 4 Vfi h! . ha . Vfi/r + 4 v2", v: + vz + Vfi + v3", where: E - narrow-sense heritability Vfi - variance among tree within plot V2 a variance of pooled error V3 - variance among male parent in", I variance among female parent within male parent. 26 4. Q9;:§1§§193,- Correlations between height and diameter were analyzed at ages 1, 2, and 7 years. Age-age correlations of phenotypic performance of height and stem diameter were analyzed at all years measured. All correlation analyses were calculated based on family means by utilizing Pearson product-moment correlation formula as follows (Snedechor 8 Cochran, 1967): [ix-i) (st-f) r1! \/ fat-X)5 Z (tr-f)2 where: r”.=coefficient of correlation §,! =observation unit X,Y =means of observation units BEEEL1£_AND_DI§QE§§IQN W. 1. finzziyal_;a§g,- Survival rate at ages 1-, 2-, 7-, and 9-years old of the families within each aspen taxon are presented in Table 6. Table 6 : Survival rate of each aspen taxon at ages 1, 2, 7 and 9 years (in percentages). Taxa no. of 1st 2nd 7th 9th 7 families year year year year harassment: x B-smdissntm 24 84 76 75 70 £.trsm212idss x (z-x resisssiana) 10 99 99 ‘71 65 B-tramaletsss 2: Emma 24 95 91 84 80 . zesrsndigentsta x Extrsmulsidss 21 ' 92 87 76 . 74 , miraculous: x Firming!” 69 98 92 87 83 . All t-oether _--__, _ M 148 __meg_ 3?- 82 - 7s _ *) Percentages were calculated from number of survive tree at the measured years divided by number of trees initially planted. At the 1st 8 2nd year old, the survival rate of the trihybrid (2.x Igulgauiana x 2. trgmulgides) was the highest (99%, 99%), followed by 2. tramplgides (98%, 92%), P. granulgidgs x R. grandidgntata (txg crosses) (95%, 91%), 2. grandidgntata x B. tngmnlgidgs (gxt crosses) (92%, 87%), then B. grandidgntatg as the lowest (84%, 76%). At the age 7 8 9 years, the survival rate of trihybrid families dropped into the lowest (71%, 65%), while the 27 other taxa remained in the same order. 2. trgmulgides was the highest (87%, 83%) followed by txg crosses (84%, 80%), gxt crosses (76%, 74%) then 2. grandigentata (75% and 70%). The drastic drop of survival rate of the trihybrid might be caused by the attack of the poplar gall beetle (fiapgzdg ingrnata). This insect was the most destructive insect in this progeny test. At two years of age, it attacked and produced galls on trihybrid trees at twice ' the rate as on bigtooth aspen (Reighard 1984). 2. fiteg_diaggter;and_hg1gh;,- After testing the homogeneity of variances among families across taxa, analysis of variances for stem diameter and for height at all ages measured were conducted. The analysis of variances showed significant differences (P<0.01) among taxa and among families within taxa at all ages (Table 7). Table 7. F value of analysis of variance of stem diameter and height . among taxa and among family within taxa. j Source Diameter Height ' 13:. 2nd 7th 9th 1st 2nd 7th year year year year year year year Rep. 7.04" 5.24“ 10. 55“ 8.07“ 6.47“ 12 .25“ 17. 54“ Taxa 10.59“ 8.66" 5.30“ 4.55“ 23.49“ 19.77“ 5.29" Family 2.28“ 1.84“ 2.18“ 2.53" 2.20“ 1.89" 2.08“ (Taxa) MS 0.0974 0.3420 2.4978 4.5410 831.10 2283.89 21194.75 Error ". Significant at P<0.01 28 29 Means stem diameter of each taxa at ages 1, 2, 7 and 9 years are presented in Table 8 and Figure 2: means for height at ages 1, 2, and 7 years are presented in Table 9 and Figure 3. At one-year-old, the mean diameter of the trihybrid (1.40) was significantly larger and different from the means of other taxa. 2. txgmglgidgs (1.35) was signi- ficantly larger than txg crosses (1.23) and B.g:andidgntata (1.07), but not significantly larger than gxt crosses (1.28). The reciprocal hybrids (txg and gxt crosses) were not significantly different from each other but significantly larger than 2. grandidgntata. 2. grandidgntatg was the smallest and significantly different from other taxa. At second year, again the trihybrid was the largest (2.40) and B. grandidgntata was the smallest (1.91). The trihybrid was significantly different from txg crosses (2.20) and 2. grandidgntgtg (1.91), but not significantly different from 2.3:emulgides (2.39) and gxt crosses (2.28). Reciprocal hybrids (gxt and txg crosses) were not significantly different from each other, but they were significantly different from E. grandidgntata. B. grandidgntata was the smallest and significantly different from other taxa. At age 7 and 9 years old, the order, based on stem diameter, was gxt cross (6.08 and 8.94) as the largest, 30 followed by txg cross (5.84 and 8.58), 2. tremuloides (5.81 and 8.46), trihybrid (5.83 and 8.40), then 2. grandidentata (4.69 and 7.14). Those four former taxa were not significantly different from each other but were significantly different from E. QIQDQIQEDLAEA- The pattern of height growth was similar to that of diameter. At the first year the trihybrid and 2.;rgmulgiggs were significantly taller than the hybrids (gxt and txg) or B. grandidentata. The hybrids (gxt and txg) were not significantly different from each other, but significantly different from B. SIADQIQQDEALQ- At the second year, 2. tzgmulgiggg was the tallest and different from other taxa. The trihybrid and gxt hybrid were second, followed by txg hybrid, then 2. grandidggtata as the lowest. At 7 years, the highest was gxt hybrid. The gxt hybrid was significantly different from the trihybrid and B. grandidgntata, but not from txg hybrid and 12. 3231119121512:- 2- madldentata was significantly shorter than other taxa. Graham et al. (1963) reported that, when growing together, 2. grandidgntatg outgrows B. trgmulgidgs. In contrast, this test showed that at all years measured 2. tzgmglgidgs always outgrew 2. grandidentata. This_ result is similar to a previous study by Brissette and Barnes (1984) of two-year-old aspens in southeastern Michigan. 31 .mo.o u scene on AOOHV en.o unencuuao Sauceoqudco«s no: one moaned ease ecu cud: use» ease on» as sense 03» wet: ..uonusoou was. 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Means of height at ages 1, 2 and 7 years. 34 A rapid juvenile growth is positively correlated with aspen survival rate (Reighard 1984, Pauley 1963, Hattemer and Seitz 1967, Morhdiek 1979). In this test, the correlation appeared only in the early growth (first year) and became obscure in the following ( 2nd, 7th and 9th ) years. At the 1st year, the fastest growing taxa was the highest in survival rate and the slowest growing taxa was i also the lowest in survival rate. At the 2nd year the relationship somewhat degraded and at the age 7 and 9 this relationship no longer held true. This phenomenon might be caused by the attack of some diseases or insects that had different severeness and preferences regarding taxa. As reported by Reighard (1984) some diseases and insects, such “mammamimandms ghprggyiatgs, were evidence at age two years. He reported that 25;;gmnlia_infected mostly trembling aspen: it was intermediate in the hybrid and less in bigtooth aspen. 5.1nggnata attacked the trihybrid twice as much as bigtooth aspen. Compared to other taxa, 1.abbrgyiatu§ mostly damaged 2.x smithii. At older age, there were also other insects and pathogens at work in this plantation. For example, after about 5 years, cancer diseases became very important. Brissete and Barnes (1984) reported that theify hybrid of 2. trgmulgidgg and 2. QIADQIQQDSAEA showed neither hybrid vigor nor marked growth inferiority compared with progeny of their parents. The hybrid of B. trgmulgidgs and 35 2. grandidgntata is intermediate in most morphological characteristics (Pauley 1963, Barnes 1961) and has a tendency to approach, but not exceed, typical rapid growth of B. txgmulgidgg in height (Pauley 1963, Einsphar 8 Benson 1964, Henry 8 Barnes 1977, Brissete 8 Barnes 1984). The performance of this progeny test showed similar results at the first- and the second-year. The height and stem diameter of E. granulgides x 2. grandidentata hybrids (reciprocal) were intermediate between progenies of either of their parents. However, at ages 7 and 9 years the results were somewhat different. The mean stem diameter and height of the hybrids exceeded, although not significantly, 2. Miniseri- Moreover, following growth of each taxa for several years (Figure 2 8 3) and number of families of each taxa that comprised 15 (10%) best families (Table 10), it seemed that dominancy of 2. tremglgideg as the fastest growing families at the early growth was replaced by the hybrids (txg and gxt crosses) at the older ages. Even though, at age 9 years, the analysis of variance still did not show significant difference (P>0.05) between E. granulgidgs and the hybrids, the mean value of stem diameter and height showed that the hybrids, which were smaller than 2. trgmglgiggs at ages 1 and 2 years, gradually became larger at ages 7 and 9 years. The dominancy of B. trgmglgidgg families in the 15 best 36 families at the first-year decreased with the time. On the other hand, the hybrid (gxt and txg crosses) families, that were less representative in the 1st year, increased along the years and became dominant at the 7th and 9th years. Table 10. Number of families of each aspen taxon that comprise 15 (10%) of the best families in height and stem diameter at ages 1, 2, 7 and 9 years. No. of Diameter Height Taxa families , planted 1st 2nd 7th 9th 1st 2nd 7th yr. Yr. Yrs yr. yr. yr. ‘ E-msdidestsfa x 24 o o o o bestow 2.5mm x 10 3 1 1 1 1.2.2: W) Edaamsbageaac z: 2 3 5 5 lbsnmEHdeMama 2. a d den x 21 2 2 5 6 lbtummdeuku Zagmmmlegmuix 69 s 9 4 3 2. m These results suggest that until age 9 years the superiority of the hybrids did not appear clearly, but indicated that hybrid vigor of the aspens' family might show up at an older age. In relation to the composition of additive and dominance variance within genetic variance (see herita- bility and components of variance page 46-56), the growth superiority of the hybrids seems to be affected by non- additive (dominant variance) genetic variance. Johnson and Larsen (cit. by Reighard 1984) also reported that non- additive genetic variance was responsible for the growth 37 superiority of hybrids between geographically isolated aspen species. The largest family mean diameters of trembling aspen, bigtooth aspen, trihybrid aspen, txg crosses and gxt crosses at age 9 years were 10.6 cm, 9.8 cm, 10.9 cm, 13.4 cm and 12.1 cm (Table 11). The tallest families at age 7 years were 875 cm, 788 cm, 826 cm, 1020 cm, and 923 cm, respectively (Table 12). These families are 25% to 56% larger than the average stem diameter and 20% to 42% taller than the average height of each taxa. They are 17% to 61% (in diameter) and 14% to 48% (in height) larger than the average of all families. The averages of the 15 (10%) best families in diameter (11.2 cm) and height (870.8 cm) were 35% and 26% larger than the average of all families, respectively. In line with the result at two-years-old reported by Reighard (1984), at 7-years-old, the early growth of these families was still comparable or greater than those reported for promising trembling aspen, hybrid aspen, triploid hybrid aspen, white poplar-bigtooth aspen hybrids and white poplar-aspen trihybrids (Pauley 1963b, Pauley 1963, Benson 8 Einsphar 1967). . .uosuouou «as no asses: asses. ussum so useeo uses—6.2.0 one: censuseoued .u .muefiquoomeou sons» some uo asses so uoeeo uoueasoaeo one: censuseouonu f Wi‘i‘iri iiuiliili' Iii |.l ‘.i .i It i. .I ll..l.lii i .1: i l l i i. ii .i. ii . l i .l l . .. .. Iiilill i 38 amnd IJ _ $2 :3 $3 $8 2.2 $2 2.: m 35 :3 .82 same amen suns some 33 jam 0 use .N Eva m6." n.m o.N odd 5.5 En min on limouJOllslll: Emu .M «new same .38 83 we: 83 an: :2 . 82 2.3 seen «2 $2 33 33 $2 magmas.“ 9: «.3 8.... Eu 22 m6 n4” e; x 33333.». 33 «mom 53 Sen :3 3: $2 2.: some $3 :2 33 some 53 as: :3 mums—squamous." «.3 ....3 8.2. 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Selection of parents is an important point for providing improved seed sources. The importance of parental selection in progeny performance of forest trees species has been recognized (Duffield 1958, Hyun 1976, Little andTrew 1976). In Massachusetts, Pauley et a1. (1963) reported that trembling aspen from the Lake states origin survived and grew better than those from Washington and Yukon territory. Progeny of European aspen (B. granulg) from central Europe grew faster than those from northern Europe (Pauley et al. 1963a). Reighard (1984), analysing all five plantations of this test at two years old, reported that the female parent showed a significant effect on progeny performance, but not the male parent. Based on this result, analysis of variance of geographic parental origin was conducted only for the female parent. The analysis showed that there was a significant difference in diameter (P<0.05) but not in height among progeny from different female parent origins (Table 13). Nine-year-old performance of this test showed that the best progenies of these aspens have their maternal parent from the central Lower Peninsula. In contrast to these; results, under greenhouse conditions, western Upper Peninsula sources of bigtooth and trembling aspen grew faster than those from other sources (Okafo, 1976). 41 Table 13. P value of analysis of variance of height and stem diameter among maternal parent origin.(Entries are means of families at each region) Source Diameter Height lst 2nd 7th 9th lst 2nd 9th year year year year year year year : Rep. 4.71" 1.92“ 3.87" 3.46“ 4.24" 4.24“ 6.10“ Taxa 7.96" 4.85“ 7.87“ 6.73" 13.83“ 9.63" 6.98” : Region 4.40" 1.95' 3.74' 2.26“‘ 1.85“‘ 2.019' I 11er 12.08“ 5.24' -S.78' 4.98‘ 2.06" 1.04" ! lower pens. :.MS Error’ 0.0287 0.1266 0.9676 246.00 '757.99 7004.29 ' . Significant at P<0.05 ". Significant at P<0.01 ‘“. Non significant at P<0.05 Southern seedlots from the same species usually grow faster than the northern ones (Wright 1976). This phenomenon is also evident in this test. Mean stem diameter of families with maternal origin from the Lower Peninsula was relatively larger than those from Upper Peninsula. At age 9 years, the mean diameter of families with maternal origin from the Lower Peninsula overall was 7% larger than those from Upper Peninsula. Duncan's Multiple Range Tests showed significant difference between central Lower Peninsula and Upper Peninsula parental origin, but not between northern &- southern Lower Peninsula and Upper Peninsulas. However, mean stem diameter and height of families from the northern, central and southern Lower Peninsula were 6%, 12% 42 and 3% larger than those from Upper Peninsula, respectively (Table 14 & Figure 4). This is consistent with (but less than) results reported by Wright (1976). He reported that trees from the Lower Peninsula grow 10% to 20% faster than those from Upper Peninsula if tested in Lower Peninsula. Although, there was no significant difference in height among families with different maternal origin, the 15 best families in height at 7 years were dominated by families that have maternal origin from the Lower Peninsula (Table 17). A similar result was also evident for stem diameter at 9 years old (Table 16). These results may indicate that for a plantation in the Lower Peninsula, a maternal parent from the Lower Peninsula, especially the central Lower Peninsula, will give a better progeny than those from the Upper Peninsula. The different growth between progenies with maternal parents from the Upper Peninsula and Lower Peninsula might be caused by several reasons. Trees from the Upper Peninsula were separated from those from the Lower Peninsula by the Strait of Mackinaw that forms a natural restriction for gene exchange. Therefore, different natural selection pressures could result in development of races that are more or less distinct (Wright 1976). Trees are genetically adapted to photoperiod of their native habitat (Spurr & Barnes 1980). The growing season in northern latitudes has longer days than the southern 43 Table 14. Mean of stem diameter of aspen families among geographic areas at ages 1, 2, 7 and 9 years. Region Latitude lst year 2nd year 7th year 9th year Lower 41.8°-—43.0°N 1.32 t 2.24 ' 5.532” 8.08“ Peninsula (1162) (1062) (1032) (1032) 43.0°-44.2°N 1.26 ' 2.23 ' 6.10 ' 8.83 ' (1112) (1062) (1142) (1122) 44.2°-4S.4°N 1.25 ' 2.27 ‘ 5.60‘h 8.37“ (1102) (1082) (1042) (1062) combined 1.28 2.25 5.76 8.43 (1122) (1072) (1072) (1072) ; Upper 45.4°-46.6°M 1.14b 2.11 ' 5.37b 7.86” 1"“‘n'?¥‘. 6(100"- (1°03)-_ -Percentages were calculated based on the mean diameter of families from Upper Peninsula. ~Any two means in the same years with the same letter are not significantly different at alpha-0.05 according to Dmcan's mltiple range test. Table 15. Mean of height of aspen families among geographic areas at l, 2 and 7 years 4 Latitude 2nd year 7th year 1st year m: 41.80-43.00" 113057 . 187e98 . 674019 . Peninsula (1062) ( 972) ( 972) 43.0°-44.2°N 108.30 2 194.91 2 724.27 ‘ (101‘) (101‘) (104‘) 44.2°-45.4°N 112.13 ' 199.45 ‘ 684.30 ' (1042) (1032) (982) combined 111.29 193.25 694.64 (103‘) (100‘) (100‘) "FPO! 45.4o-46.5°N 107e58 . 192e98 . 697.00 . - Pgnin’“1‘ (10°11_ _. ,(}993’um _ ({99§1_- -Percentages were calculated based on the mean height of families from Upper Peninsula. -Any two means in the same years with the same letter are not significantly different at alpha=0.05 according to Duncan's multiple range test. 41) cm 10 9 8 7 6 5 4 3 2 . 1 0 tat-year 2nd-year 7th-year 9th-year - g 41.6 - 43.0 N_ (LP) : 44.2 - 46.4 N_ (LP) : 4&0 - 44.2 N (LP) : 46.4 - 40.0 N.(UP) note : any two means in the sales year with the same letter are not slgnlfloantiy different at ”0.06 aooordlng to Duncan's multiple range test Figure 4. Means of stem diameter among geographic areas at ages 1. 2. 7 and9 years cm 800 600 400 200 - , °_ :: (\\ .. 1st-year 2nd-year 7th-year - :41.8 - 46.0 N _(LP) 844.2 - 46.4 N (LP) , 46.0 - 44.2 N,(LP) - ,46.4 - 46.6 N (UP) note 2 any two Ileana In the same year with the sales letter are not signifioantly different at we.“ sooordlng to Duncan's Iaultlple range t." Figure 5. Means of height among geographic areas at ages 1. 2. and 7 years. 45 Table 16. The fifteen best aspen families (in diameter) at age 9 year. Parental origin Accession Diameter number (cm) Maternal parent Paternal parent County Region County Region 70071' 10.25 Montcalm II Sanilac II 2901054 10. 39 Lake 11 Chippewa 1v 90038d 10.53 Lake II *) open pollination *70057' 10.71 Calhoun I Iron IV 70033c 10.77 Marquette 1v Oakland I 70070' 10.77 Moncalm II Van Buren I 60001‘ 10.87 Calhoun I Alpena III *90125‘ 10.69 Wexford n Ralkaska III *70044° 11.13 Roscommon II Oakland I 70079“ 11.17 Van Buren I Oscoda III *70081' 11.31 Wexford II Benzie III *70024° 12.03 Iosco III Gladwin II *70078‘ 12.07 Van Buren I Iron IV *70004° 12.41 Branch I Clare II *70043° 13.43 Roscommon II Ingham I . I _m-_11“,,11171_, )===_, ..slso excellent in height at 7 years 22.2mm x (3.: W) clkflumflaflhelrzessflnEEQEa 13.393161“; x 2.3mm 0.2mm 8 2mm 46 Table 17. The fifteen best aspen families (in height) at age 7 year. 1-- _______.__=.______..=__ Is , Parental origin ii-l ) Accession Height 1 numbgr (cm) Maternal parent Paternal parent ; County Region County) Region ; 600041’ 616.42 Calhoun I Iron 1v i 60006h 626 . 33 Calhoun I Oscoda III 70005° 831.63 Branch I Marquette IV 70071“ 835.67 Montcalm II Sanilac II *70057‘ 836.90 Calhoun I Iron 1v 90117)I 645 .92 Oceans 11 Huron n «90105‘l 851.80 Lake 11 Chippewa 1v *70081‘ 852.20 Wexford II Benzie III . 90106d 653 . 25 Lake II Brunet 111 W *90125‘ 874.92 Wexford II Kalkaska III : *70044° 876.60 Roscommon II Oakland I ; *70004° 887.00 Branch I Clare II } *70078‘ 923.50 Van Buren I Iron Iv *70024° 928.67 Iosco III Cladwin II *70043‘ 1019.75 Roecommon II Ingham I i * .a1a0 excellent in stem diameter at 9 years liftmsmhuaulxtf xrmuaednu) - aumiLemuas 47 latitudes. Therefore, if planted in southern latitudes, northern trees will stop growing sooner than southern trees due to the shorter daylength. Reduced daylength will trigger growth cessation (Vaartaja 1960). Trees also genetically adapted to the temperature regime of their native habitat (Perry 1962). Brissete and Barnes' (1984) reported that aspen from lower summer daily mean temperature habitats exhibit low rates of photosynthesis and high rates of respiration, when preceded by higher daily mean temperature. Therefore, when planted in the Lower Peninsula, aspen progeny from the Upper Peninsula may grow slower than those from the Lower Peninsula. Summer daily mean temperature at high latitude habitats is lower than those at lower latitude. In agreement with the result reported by Reighard (1984) from the same plantation at age two years, but with different sample, analysis of variance using the nested design in Table 3 showed that the variance component of female-within-male was significant (P<0.05), but not the male component (Table 18). The component of variation associated with male parent, female-within-male parent, error pooled and trees- within-plot of stem diameter and height for all years measured are presented in Table 19 and Figure 6 a 7. 48 Table 18. P-value of ANOVA derived from the nested design (NC design 1) at Table 3. Diameter l Height ; Sources 1 1st 2nd 7th 9th 1st 2nd 7th 1 year, year year year year year year Z Rep. 4.05“ 1.88“ 5.92" 3.06' 2.65“ 2.25" 6.73“ i . Male 1.67“ 1.83” 1.21" 1.04“ 2.22“ 1.90" 1.11” Female 2.32“ 2.16“ 2.99” 3.11“ 2.65” 2.44“ 3.09“ ) (male) . ER”: 0.093 0.310 .2._ 376 4. 37 .05 1954.06 _ 1832.95 :2 Significantly different at alpha level -0.05 ) Significantly different at alpha level -0.01 ‘“) Non significantly different at alpha level -0.05 Male variances of stem diameter and height were small and not significant at all years measured. The trend of male variance was decreasing along years. It ranged from 8% at age two years to 1% at age nine years for diameter, and from 12% at first year to 1% at age seven years for height. On the other hand, female variance was relatively larger than male variance and significant at all years measured. The female variance seemed to increase with the age. It ranged from 10% at the first year to 15% at age nine years (for diameter), and from 13% and 11% at the first- and second-year to 16% at age seven years (for height). These results showed that variation in progeny of aspen were affected more by the female parent than by the male parent. It also indicates the importance of selecting female parents in a mating design. However, Other research ? i 49 ..uossceouu .ue> use ..uosoom. uouuu.we> .AOAeSVOHeeou .ue> .ese8.we> no seuou on» so useen one: eoueusoowom \s same. seem. seen. same. sash. soon. shew. .uoss. oo.eepos mm.~m>s mm.nee smh.m neo.n new. nos. eons .ue> sese. same. .oon. seen. .aen. same. .aoe. .eesoos. ma.-nms mo.vmms mo.eo> hon.e can.“ csn. «co. uouwo .we> .aosc sass. .ans. .ems. .ans. sees. .aes. .eses. ms.mnnh so.m~m mo.on~ moh.s use. moo. «no. oseeou .we> .os . .ee. seas. .es. sun . .ao . see . mm.eso ms.ooe m~.~«~ mmo. «es. «no. mso. osee .we> wee» can seek us« see» you wee» sum week sub ween usN ueoa.uen \susmsom \euouoeess eoowsom .smseou ossuee s 02 sowu uo>swou senses use wouoeesu sous no 00seswe> uo ausosomeoo .ma Canes 50 60% 40% ’ 202 — 02 ' . . . . age in year 8 percentage was calculated from total ail variances -— Var. msie ‘i— Var. female (male) + Var. error (pooled) ‘9' Var. tree (plot) Figure 8. Components of variance of stem diameter at ages 1. 2. 7 and 9 years 50$ 40% 30" 20% 10" 0'4 derived from NC 1 mating design (in percentage). 42‘ 41‘ F .1" age in year $ 90'0““... '8. calculated from total Iii variances + Var. female (male) "9- Var. tree (plot) ""- Var. male + Var. error (pooled) Figure 7. Components of variance of height at ages 1. 2. and 7 years derived from NC 1 mating design (in percentage). 51 by Mohrdiek (1979), who crossed 2. tremulg with B. granulgiggg, indicated that selection of either male or female parent was very important. The non-significance of the male parent in this test could be caused by the strong effect of maternal parent or by experimental error (Reighard 1984). The variance caused by experimental error were relatively high. It ranged from 82% to 86% for diameter, and 75% to 83% for height. A further test by using reciprocal parents (male parent nested to female parent) might be worthwhile. Assuming that there was no epistasis in the genetic variance, the estimate of additive and dominance component variances and the narrow sense heritability were calculated and presented in Table 20 and Figure 8 to 11. Heritability estimates apply only to a particular population, in a particular environment and in a particular point in time (Zobel and Talbert 1984). Einsphar et al.(1967) reported narrow-sense heritabilities of 0.24 (height) and 0.35 (diameter) for full-sib families of trembling aspen. Reighard (1984), using the same plantation of this test with a different set of samples, found narrow— sense heritabilities of 0.31 (height) and 0.39 (diameter) at the second year. In this test, narrow-sense family_based heritabilities for diameter and height were high at the first year (0.50 for diameter, 0.83 for height) and second years (0.63 for diameter, 0.66 for height), but it changed 52 ..smmsv eusewsx use wosessem 0» ossuwoooe essfiwou assess an uouessuseo owe: assasssneuswoz \« .30s500wu .we> use ..u0soomvwowwn.we> .oswosom.we> uo «once as» so uooeo owe: someusoowom \s ~-iii.’l. iiiii- ii .. .. ...... emo. com. «me. eso. Coo. men. one. oowuuosuesh coo. nmo. one. one. ems. «no. new. 2s ocean assess M smsssmemsummuumoumummuueu . seen. seems same. .anno Asses seems seems ‘ om.oeeos mm.~mes em.noo see.» neo.n ecu. nos. suose. eons seem. soon. sesn. seen. some. .oen. .ann. soosoos. r mo.-nos oo.emos mo.eoe eon.e oen.« osn. eeo. nouns L umeequewlaeueusemuqnsu save. seems sase. swseo seen. «was. sesnc so + es 1 oo.~smeu oe.oss~ ~s.eee ems.» emo.n use. one. osuoooe , .sov. see e sea . .eoe. some. see c sons. .6. m~.vmoau mo.nee «o.om mum.o ooo.n moo. eno. oooeosaoo see . .ooa. some. .as . .am . some. sees. 2e. . sn.omo~ em.e~os os.eoo new. men. new. «no. oesuseoe L «moasuewlmquummm wee» sup wee” usn wee» was week 53 weox sub weoa usn weoN was _. umuuflom M \susOsom \swouoaess 1 .oosoeo mosses s 82 soon ooesuoo ”SOdOfl Ufld UGUOBUdU ECU. U0 M$dddfldflflh0§ OIQCII3OHHIC USU OOGQdHI? HGHGQBGOHA>GO ”Gd OOAUOGOO .ON OHQGH 53 80" 60% " us 41‘ 40% 20$ 02 ‘ 1 2 ' 3 4 5 8 7 8 9 age in year 6 percentage was calculated from total all variances (sodoe) -‘- Var. additive is) —+— Var. dominance (d) + Var. genetic (sec) ‘9' Var. environment (a) Figure 8. Genetic and environmental variance estimates of stem diameter at ages 1. 2. 7 and 9 years (in percentage). us see 00* see ~—e 44s «2 462 40* see ass ass 20". - as as 42 0“ I 1 l 1 1 __1 1 2 3 4 5 8 7 age in year 7. percentage was calculated from total all variances laOdoe) f-r- Var. additive is) -*- Var. dominance (d) ‘4‘" Var. genetic (rd) “'9' Var. environment (a) Figure 9. Genetic and environmental variance estimates of height at ages 1. 2. and 7 years (in percentage). ' 54 0.7 F 0.6 ' 0-5 ..... 0.4 r 0.800 0.3 r 0.2:. 0.228 0.7.. 0.1 ' 6.6a 0.020 0 l ' I i 1 1 , .an 1 2 3 4 5 6 7 6 9 age in year I - --. mass" . +. ":.:‘.:::::'.'.:'.'“ Figure 10. Family- and individual tree-based heritability of stem diameter at ages 1. 2. 7 and 9 years. age in year fatally-based individual tree-based , heritability , -8- , heritability . Figure 11. Family- and individual tree-based heritability of height at ages 1. 2 and 7 years. 55 drastically at 7 years (0.14 for diameter, 0.07 for height) and at 9 years (0.03 for diameter). A similar result also showed up for single-tree based heritability, but all single-tree based heritabilities were smaller than those based on family. The decrease of heritabilities might be caused by the change of composition of additive and dominant variance within genetic variance, and by environmental effects. As trees mature, the heritability changed markedly due to environmental change and composition change in genetic control of the characteristics (Zobel and Talbert 1981). In this test, the genetic control (genetic variance) of stem diameter was relatively high and slightly increased with ages (from 31% in the first year to 41% at nine years old). For height, the genetic variances were also high, moving from 41% at the first year to 36% at the second year and then up to 44% at 7 years. The composition of additive and dominance variances within genetic variance also changed with the age. The additive variance component dominated genetic variance at the first and second year but decreased markedly with age (from 18% at the first year to 1% at 9 years for diameter, and 39% at first year to 4% at 7 years for height). On the other hand dominance variance components that were less pronounced in the first and second years increased with the age (from 13% at the first year to 40% at 9 years for diameter, and from 2% at the 56 first year to 40% at 7 years for height). Since narrow- sense heritability is based on additive variance, the decrease in additive variance decreased the heritability. The decline of additive variance with ages often occurs in forest trees species (Franklin 1979). Namkoong and Conkle (1976) showed a marked decrease of additive variance (in height) between ages 5 and 7 years in half- sib families of Ponderosa pine. Gill (1987) reported a decline of additive variance between height at 8 or 11 years and 22 years in half-sib families of white spruce. This problem should be a major concern for a tree breeding program, because if there is no additive variance at a certain time there will not be any prospect for improving general combining ability by doing selection at those times (Wright and Talbert 1984). The declining of additive variance in this test might be caused by the interaction between inter-tree competition and expression of genetic variance. Franklin (1979) mentioned that inter-tree competition is a major causal factor in the behavior of additive genetic variance when the stand is developing. The increased growth of trees may increase inter-tree competition, influencing the additive genetic variance. Cannell (1982) also showed evidence, in open pollinated Sitka spruce, that the decline of additive variance was caused by the increasing inter-tree 57 competition. However, the phenomenon of interaction between competition and genetics has not yet been widely studied in forest tree breeding. The contribution of components of variance containing non-genetic factors (var.error and var.trees-within-plot) to the total variation was relatively large (ranging from 69% to 58% for diameter and 64% to 56% for height). This large non-genetic variance may indicate that silvicultural practices for increasing growth is important. It should be noted that these components of variance were derived from a specific population involving a mixture of inter- and intra-specific crosses. An analysis based on inter- or intra-specific crosses separately might give a different result. Correlations 1. W.- The coefficient of correlations between stem diameter and height at ages 1, 2, and 7 years, for all taxa together and for each taxa separately, showed that stem diameter and height were significantly correlated (Table 21 & Fig 12). The coefficient of correlation ranged from 0.88 to 0.90 for all taxa together, and from 0.73 to 0.97 for_each taxa separately, except for 2.;zgmulgiges x £.grgng1gggtata at the first year (0.44). Reighard (1984), analyzing all five plantations in Michigan at two years old, found a 58 Table 21. Coefficient of correlation between stem diameter and height at ages 1, 2, and 7 years. 1:1— lst year 2nd year 7th ’ year 0.83" 0.86" 0.90" 0.89" 0.97” 0.73" 0.44" 0.79” .89” 0.73" 0.90” 0.87” 0.82" .84" 0.89” 0.88” ”) Significantly correlated at alpha - 0.01. range of coefficient correlations from 0.73 to 0.89. Another study on Leuce progenies by Mohrdiek (1979) also showed that height and stem diameter were highly correlated. The high correlation between stem diameter and height is an advantage for a tree improvement program. It gives a possibility to estimate height by only measuring stem diameter, or vice versa. M\\\\\\\\\\\\\\\\\\\swm .60 2. Agg:ggg_gg;;§1g§123‘: Age-age correlation of a trait is a principal tool in forest genetics for calculating the gain from juvenile selection to a future breeding program (Zobel and Talbert 1984, Namkoong 1979). Age-age correlations are influenced by growth rate (Namkoong and Conkle 1976), site, stocking and competition (Franklin 1979). The age-age coefficients of correlation of family-mean diameter between ages 1, 2, 7, and 9 year, and those of height between ages 1, 2, and 7 years are presented in Table 22. The correlation of phenotypic performances between ages 9, 7, 2, years and age 1 year (for diameter) and between ages 7 & 2, years and age 1 year (for height) of all taxa combined as a group, were significance (P<0.05). However, the coefficient of correlation gradually decreased along with the distance between ages measured (Figure 13). Based on age 9 years for diameter, the coefficients of correlation for ages 7, 2, and 1 were 0.94, 0.66 and 0.54 respectively. Based on 7 years for height, the coefficients of correlation for ages 2 and 1 were 0.62 and 0.52 respectively. These results are similar to those from Leuce progeny reported by Mohrdiek (1984). He found numbers of 0.952, 0.934, 0.828, 0.554, 0.483 and 0.462 for correlation between age 25 to 15, 11, 9, 3, 2, and 1, respectively. Reighard (1984), found a coefficient 61 correlation of 0.48 between one-year-old trees in the nursery and 2-year-old trees in the field. The significance of the correlation may give a possibility to do an indirect selection, based on height or diameter, earlier than 9 years old. Analysis for each taxa, separately, showed that families of bigtooth aspen, trembling aspen and the B. trgmulgides x B. grangiggntgta hybrid had significant correlations for all ages, but not for the trihybrid and the 2. gzgngiggntata x 2. tzgmglgiggg hybrid. This may indicate that, if early selection will be done based on families of each taxa separately, it will only be partially effective for bigtooth aspen, trembling aspen and 2. trgmulgiggg x R. grandidgntgtg, but not for the trihybrid and hybrid 2. grgngidgntgta xyz. tzgmglgiggs. 62 mo.ovm we useosussmso :02 .ns soo.ove ue oceosuscosm ... mo.ove ue sooosesomsm .. .Mxeu sse umssneoo .m mous0sseowu.m x mousOsseowu.M .0 emusOHsEmwu.M x mmmmmmmflmmmmm.m .u mmmmmmmwmmmmm.m x nousOssEOwu.M .0 esesseessowx.m w nousossemwu.m .n .mmmmmmuwmqmmm.m x mmmmmmmflmdmmm mmdmmmm .e a ouos a s .oeo_¢ n» :exo.c :eno.. 668:. . :eno.. . . . . .e :mno.o s :~o o a :~m o c e a z :ee.o .6 abno.h :exo.. :mno.. _ :mno.c :3ao.c 238:” .fiwgw.w .:mmo.a :sno.. .. . . .sto.o :euo.o ..on O n . No.0 .0 ’o.° .0 I I. C O . O. .. .- ....m o s :3 o c N o .. :2... .a some .o :exo.. gene 3. _ :sso.c Noszo . .. . .. ............. “Mm“ s. .wwn.6 :qeo.. :swo.. :xso.. ~m.o .6 :oo.o .6 :o..o .o .¢o~.o .o :~o.o .o «as.o .o :m~.o .o .~o.o .o :am.o .o :oo.o .o :os.o .. .o2.o .. . s.oa= ges.o .o :6..o .a :~o.o .o W : :ee.e .. :~o.o .. :o~.o .. s m s.Ea_a o e a e a. z N a z s so: e.a._a 5....6 ~.e.1a ..ea_a .mLoma a com a .N .s some moose s=m_o: Coo Lososo_u seam co oossmsoocoo co somsuwccoou o_o»sooo;o .- seams QQEQLUSION The results of this study indicate that the hybrids between E. trgmulgiggs and 2. grgngiggntgt; were able to establish themselves as readily as either 2. gzgngiggntgtg or 2. tzgmglgiggg. For the families used in the experiment, the survival rate of the hybrids and the trihybrid, especially at the first and second year (87-99%), were high and almost equal to the progeny of their parent. However, the survival rate of the trihybrid at 7 and 9 years of age (71% and 65%) as slightly less than the progeny of either 2- stabilisation or 2- M12193- 2. gzgngidgntata was the least competitive among the taxa investigated. Height and stem diameter growth of the hybrids were initially intermediate between B. trgmglgiggg and B. grgggiggnt§t_. However, at ages 7 and 9 years the hybrids surpassed either of their parents. The hybrids, which were initially not dominant among the 10% best families, became dominant at ages 7 and 9 years. These results indicated that hybrid vigor might exist in aspen families. The superiority of the hybrids might not appear early on, but rather show up in lates years. Further study of this test at an older age might be worthwhile. 64 65 Variation of the progeny was affected more by female parent than by male parent. The male parent variance was small, not significant, and decreased with age. On the other hand, the female parent variances were significant and tended to increase with age. The performance of the progeny was also affected by geographical origin of the female parent, in term of stem diameter but not height. For plantations in the Lower Peninsula, female parent from the Lower Peninsula had better progeny than those from the Upper Peninsula. Among female parents from the Lower Peninsula regions, those from the central Lower Peninsula generated better progeny. Narrow-sense heritability estimates of stem diameter and height traits were relatively high in the early growth but decreased extremely with age. The family-based heritability was relatively high at the first-year (50% for diameter and 83% for height) but declined to below 10% at 7 years (for height) and 9 years (for diameter). The individual tree-based heritability had a similar pattern and was always smaller than the family-based heritability. The proportion of genetic variances for stem diameter (31%-41%) and height (36%-44%) was relatively large and slightly increased with age. This is an advantage for tree improvement program. However, composition of additive and dominance variance within the genetic variance changed with time. The additive variance decreased along the years 66 and was very small at ages 7 years and 9 years. The dominance variance increased with the ages, and at age 7 years and 9 years became the major proportion of the genetic variance. The proportion of non-genetic variance (environmental variance) was relatively high (range from 69% to 56%) and slightly decreased with age. Stem diameter and height traits were highly correlated to each other. The coefficien of correlation for all taxa, combined, ranged from 0.88 to 0.90. Age-age correlation among ages 1, 2, 7 and 9 years (for stem diameter) and ages 1, 2 and 7 years (for height) were also significantly evident for all taxa together. These results provide some justification for doing indirect selection at an early age. BBQQHHEEDLIIQEE The objective of a tree improvement program is the development of improved trees and mass production of improved seed or propagules at any stage of their development, for immediate need (Zobel and Talbert 1984). The progeny test in this experiment showed that hybrid vigor seemed to appear in the F1 hybrids of trembling aspen x bigtooth aspen (reciprocal). It means that, for plantation in southern Lower Peninsula of Michigan, production of this hybrid could be more beneficial. However, since the superiority of the hybrids seemed to be controlled by non-additive genetic variance, the F2 progeny of the best F1 hybrids could not be guaranteed to be better than the F1 itself, they might even be worse (Wright, 1962). Based on these reasons, mass production of selected F1 hybrids by clonal means or other plantings of parental species will be better than production of untested F2 hybrids from the best F1 hybrids. Mass production of the selected F1 hybrids may be accomplished through several methods. 67 68 In aspen, vegetative propagation of the F1 hybrids is a possibility. Aspen is noted for its ability to regenerate vegetatively by adventitious shoots or suckers that arise on its long lateral roots (Debyle and Winokur 19..). An average of two suckers can be produced from 2.5 lineal centimeters of 0.63 cm to 1.27 cm diameter root cuttings (Schier and Campbell 1980). Vegetative propagation also has the advantages of perpetuating preferred genotypes. Another way to produce selected F1 hybrids is by establishing seed orchard that consists of parental trees of the selected F1 hybrids, followed by controlled pollination of specified parental combinations. Parental trees should be collected clonally, while pollination can be done by using the cut-branch method (Einsphar and Benson 1964), or by using a wind-pollination method. By manipulating trees arrangement in a specific design, pollination can be directed for specific parents. The cut-branch method has generally been used in artificial seed production of aspen hybrids. This method may produce a large number of seeds. Benson (1972) reported that 700 seeds per catkin can be produced with this method, but the average production ranged from 150 to 300 seeds per catkin. The cut-branch method is also easy to handle and gives a guarantee in producing pure seed from specified parental combination. Another advantage in using cut-branch method is that we do not need to design any specific lay 69 out for parental trees in the seed orchard. However, this method might need a high capital input for collecting catkin-bearing branches and for other expenses. When wind-pollination is a preference, a chessboard distribution design is recommended (Klaehn 1960, Giertych 1975). This design simply alternates two selected clones (one pair of parents of a selected F1 hybrid) in each row and column of the orchard. To avoid intra-specific hybridization, only one pair of parents (two clones) is allowed in an orchard. Since aspen is a dioecious species, selfing will not be a problem. When more than one pair of selected parents are needed, a group of small orchards in which each orchard has difference pairs of parent might be an alternative. In this case, the distance between orchards should be far enough, or the orchards should be isolated to each other, such that unwanted crossing can be avoided. Since aspen is prolific and produces tremendous seeds per individual seed-bearing female (Graham, et al. 1963), a small number of selected female parents at each orchard may produce a huge number of improved seeds. The wind-pollination method offers the simplest and cheapest route to production F1 hybrids seed production. However, since the flowering time of trembling aspen and bigtooth aspen is slightly different, this method must be recommended with caution. 70 Another alternative for producing improved seed in the next generation is by testing the F2 hybrids. Selected F1 hybrids can be crossed to each other such that a sufficient wide genetic base of F2 hybrids can be made. A progeny test on the F2 hybrids could be done at an early year. By this time good combining F1 hybrid parents can be identified. If the genetic variance is dominated by non-additive components, a similar method for producing F1 hybrids can be applied for producing improved F2 hybrids. If the genetic variance is dominated by additive components, clonal seed orchards from F2 hybrid can be established. While producing improved seed for short-term objectives, continuous improvement program can be done by simultaneously increasing the additive genetic variance in the parents (based on the F1 Hybrids) that have exhibited the best specific combining ability and then mating these improved parents for subsequent generations of F1 hybrids. Namkoong (1979) suggested that direct recurrent selection based on general combining ability would be easier and just as effective as method based on specific combining ability. Continuous observation and evaluation of phenotypic performance and the behavior of the genetic control of growth traits at older years would be valuable. Concurrently, improvement on other traits such as pest and diseases resistance, stem-straightness, and flowering time should also be done. LI§I_Q£_B£ZEB£££EE Andrejak, G.E and B.V Barnes, 1969. A Seedling Population of Aspens in Southeastern Michigan. Mich. Bot. 8:189-202. Barnes, B.V, 1961. Hybrid aspens in lower peninsula Michigan. Rhodora 63(775):311-324. Barnes, B.V, 1969. Natural Variation and Delineation of Clone of 2929199.tremuleides and Eigrandidsntata_in Northern Lower Michigan. Silv. Genet. 18:130-142. Barnes, B.V, 1978. Morphological variation of families of trembling aspen in southeastern Michigan. Mich. Bot. 17(4):141-153. Barnes, B.V, 1975. Phenotypic variation of trembling aspen in western North America. Forest Science 21:319-328. Becker, W A, 1984. Manual of Quantitative Genetic. Fourth edition. Acad. Enterprises,'Pullman, Washington. Benson, M.K. and D.W. Einspahr. Early Groeth of Diploid, Triploid and Triploid Hybrid Aspen. Forest Science 13(2):150-155 Benson, M.K and G Dubey, 1972. Aspen Seedling Production in a Commercial Nursery. Inst. Pap. Chem. Genet. & Physio. Notes No.12, 7p. Blyth, J.E and W.B Smith, 1982. Pulp Production in Lake State by Counties, 1981. North Central For. Expt. Sta. Res. Note NC-287, 4p. Brissette, J.C and B.V Barnes, 1984. 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A rapid method of vegetative propagation of aspen and their hybrid. Forestry Chronicle 47:36-39. APPENDICES 77 four replications. Families Counties origin Table A1. Families that at age 9 years represent at least at Accession number Maternal Parent Paternal Parent 10001 Allegan open-pollination 10004 Branch open-pollination 10009 Clare open-pollination 10014 Ingham open-pollination 10015 Ingham open-pollination 10034 Ogemaw open-pollination 10037 Oscoda open-pollination 10045 Van Buren open-pollination 10048 Branch Ingham 10049 Branch Calhoun 10050 Branch Sanilac 10051 Branch Marquette 10052 Calhoun Ingham 10054 Calhoun Midland 10055 Clare Oakland 10056 Ingham Chippewa 10057 Ingham Clare 10058 Ingham Sanilac 10060 Ingham Marquette 10062 Iosco Gladwin 10069 Ogemaw Ontonagon 10070 Ogemaw Marquette 10071 Saginaw Oakland 10072 Saginaw Missaukee 60001 60003 60004 60006 60007 60008 60010 60011 60012 60015 Calhoun Calhoun Calhoun Calhoun Calhoun Calhoun Ingham Ingham Marquette ROBCOMOD Alpena Ingham Iron Marquette Midland Oscoda Isabella Isabella Isabella Isabella cont'd. table A1 (cont'd.) 78 Maternal Parent Paternal Parent 70004 Branch Clare 70005 Branch Marquette 70008 Chippewa Kalkaska 70012 Gladwin Chippewa 70013 Gladwin Calhoun 70016 Ingham Chippewa 70017 Ingham Midland 70020 Ingham Roscommon 70021 Ingham Roscommon 70022 Iosco Branch 70024 Iosco Gladwin 70027 Lake Marquette 70028 Marquette Ingham 70029 Marquette Clare 70030 Marquette Oakland 70031 Marquette Marquette 70032 Marquette Clare 70033 Marquette Oakland 70035 Oceana Clare 70038 Oceana Sanilac 70039 Ontonagon Clare 70043 Roscommon Ingham 70044 Roscommon Oakland 70045 Washtenaw Otsego 70047 Allegan Huron 70048 Branch Presque Isle 70050 Branch Oscoda 70051 Branch Mackinac 70052 Branch Ingham 70054 Branch Marquette 70056 Calhound Wexford 70057 Calhound Iron 70059 Clare Kalkaska 70060 Ingham Sanilac 70062 Ingham Midland 70069 Marquette Marquette 70070 Montcalm VanBuren 70071 Montcalm Sanilac 70072 Montcalm Alpena 70074 Saginaw Huron 70075 Saginaw Alpena 70076 Saginaw Chippewa 70078 vanBuren Iron 70079 vanBuren Oscoda Wexford Benzie cont 79 table A1 (cont'd). Maternal parent Paternalparent 90001 90003 90004 90005 90010 90011 90013 90014 90016 90019 90020 90021 90026 90028 90029 90037 90038 90039 90043 90044 90045 90051 90053 90054 90056 90057 90059 90063 90065 90066 90067 90068 90076 90078 90079 90082 90083 90084 90085 Allegan Alpena Alpena Alpena Branch Branch Chippewa Chippewa Clare Gladwin Gladwin Gladwin Ingham Ingham Ingham Lake Lake Luce Marquette Marquette Marquette Missauke Montmorency Montmorency Oakland Oakland Oceana Ogemaw Oceola Oceola Osceola Oscoda Wexford Allegan Allegan Branch Branch Branch Chippewa Chippewa Gladwin Gladwin Ingham Iron open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination open-pollination Gladwin Manistee Ingham Ingham Emmet Huron Oscoda Gladwin Luce Sanilac Iron cont'd. table A1 (cont'd) 80 Maternal Parent Paternal Parent 90094 90095 90096 90097 90098 90099 90100 90101 90103 90104 90105 90106 90107 90106 90109 90114 90116 90117 90118 90119 90120 90121 I 90123 : 90124 i 90125 Ingham Ingham Ingham Ingham Ingham Iosco Lake Lake Lake Lake Lake Lake Marquette Marquette Mecosta Oceana Oceana Oceana Ontonagon Ontonagon Osceola Osceola Roscommon Wexford wexford Montcalm Marquette Mackinac Washtenaw Emmet Gladwin VanBuren Benzie Emmet Alpena Chippewa Emmet Ingham Ingham Montmorency Iron Marquette Huron Wexford Iron Sanilac Midland Marquette Midland Kalkaska 81 Table A2. Families that were used for constructed NC-l mating design. Accession Number Male parent_ (Female parent