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Breeding and Genetic Resources of Five-Needle Pines

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Breeding and Genetic Resources of Five-Needle Pines Estimation of Heritabilities and Clonal Contribution Based on the Flowering Assessment in Two Clone Banks of Pinus koraiensis Sieb. et Zucc. Wan-Yong Choi Kyu-Suk Kang Sang-Urk Han Seong-Doo Hur Abstract—Reproductive characteristics of 161 Korean pine (Pinus the northeastern part of Eurasia. It usually occurs as a koraiensis Sieb. et Zucc.) clones were surveyed at two clone banks for mixed forest stand consisting of various broad-leaved tree 3 years. These clone banks were established at Yongin and Chunchon species and other conifers. Korean pine has been widely (mid-Korea) in 1983. Characteristics in female and male strobili planted as a pure stand accounting for about 30 percent of were spatial (between locations) and temporal (among investigated the yearly planting areas in Korea due to its high-quality times) variables. Broad sense heritabilities were found to vary timber and edible seeds. A breeding program for this species between 0.20 - 0.46 in females and between 0.34 to 0.56 in males. has been conducted since 1959 and has resulted in the Among 161 clones, 32 clones (20 percent of the total clones) ac- establishment of 98 ha seed orchards (Mirov 1967, Chun counted for 42 to 54 percent of clonal contribution in female strobili 1992, Choi 1993, Wang 2001). and 83 to 96 percent in male strobili, suggesting that the clonal The main goal for seed orchards is large-scale production contribution for male parents was severely unbalanced compared to of genetically improved seeds that maintain genetic diver- that for female parents. The effective population numbers varied sity to prevent inbreeding depression. Thus, the mainte- depending on time (year), location and sex. The mean values of nance of random mating among clones is one of the key relative effective population numbers at gamete levels were 0.56 in elements to successful management of seed orchards (Roberds females and 0.09 in males, respectively, and that value at the clonal and others 1991, Chaisursri and El-Kassaby 1993, Matziris level was 0.27 (0.25 at Yongin and 0.29 at Chunchon). The degree of 1993, El-Kassaby and Cook 1994, Burczyk and Chalupka 1997, Han and others 1999). sexual asymmetry (As) varied with a range of 0.03 to 0.24 at Chunchon and 0.07 to 0.44 at Yongin. The pattern of gamete The clonal contribution to seed production in a seed production within clones was highly asymmetrical as compared to orchard is one of the most important factors; genetic compo- that of other conifers. This indicates that P. koraiensis is extremely sition of the seed produced is determined by the contribu- low in male gamete production compared to female gamete produc- tions of each clone. Differences in clonal contribution have tion. been previously reported in several studies and have been attributed to genetic rather than environmental factors (Griffin 1982, Schmidtling 1983, Askew 1988, Brunet and Key words: Pinus koraiensis, strobili, clone bank, clonal Charlesworth 1995, Kjaer 1996, Han and others 1999, contribution, broad sense heritability, effective Nikkanen and Ruotsalainen 2000). population number, sexual asymmetry To date, numerous studies have been conducted to obtain information related to reproductive processes such as flow- ering characteristics, clonal contribution, and sexual asym- Introduction ____________________ metry in seed orchards. Clonal contribution within a seed orchard is commonly depicted by a flowering or cone yield Korean pine (Pinus koraiensis Sieb. et Zucc.) is a five- curve. In this method, the clones are ranked from high to low needle pine (Pinus subgenus Strobus) belonging to subsec- in flower production, and cumulative contribution (in per- tion Cembrae. The species has a wide natural distribution in cent) is plotted against the proportion of the clones. Addi- tionally, the concept of effective population number has been recently applied to the estimation of clonal contribution (Griffin 1982, Kjaer 1996, Choi and others 1999, Han and In: Sniezko, Richard A.; Samman, Safiya; Schlarbaum, Scott E.; Kriebel, others 2001a, 2001b, Kang 2001). Howard B., eds. 2004. Breeding and genetic resources of five-needle pines: growth, adaptability and pest resistance; 2001 July 23–27; Medford, OR, Our major interest in this study is to quantify the repro- USA. IUFRO Working Party 2.02.15. Proceedings RMRS-P-32. Fort Collins, ductive processes using empirical data from two Pinus CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Re- search Station. koraiensis clone banks, to survey the differences of clonal Wan-Yong Choi, Kyu-Suk Kang, and Sange-Urk Han are with the Tree contribution by means of flowering assessments, and to Breeding Division, Korea Forest Research Institute, 44-3 Omokchun, Suwon monitor the genetic diversity measured by effective popula- 441-350, Republic of Korea. Seong-Doo Hur is with the Sobu Forest Experi- ment Station, Korea Forest Research Institute, 670-4 Sangmo, Chungjoo 380- tion sizes. These include estimating heritability, gamete 940, Republic of Korea. Correspondence should be addressed to e-mail contribution, and sexual asymmetry. [email protected]. 172 USDA Forest Service Proceedings RMRS-P-32. 2004 Estimation of Heritabilities and Clonal Contribution Based on the Flowering Assessment in Two Clone Banks… Choi, Kang, Han, and Hur Materials and Methods ___________ where ai and gi are the proportions of ith clone of which male and female strobili contribute to the whole population. A Reproductive characteristics such as number of male and high maleness index of a clone indicates that the clone is female strobili were surveyed in the two clone banks of P. contributing more as a paternal, rather than maternal koraiensis. The two clone banks were established at Chunchon parent. (lat. 37∞55', long. 120∞46') and Yongin (lat. 37∞30', long. The effective population numbers at gamete level (Eq. 1 127∞20') in 1983. A total of 167 clones were grafted at and Eq. 2) and clonal level (Eq. 3 and Eq. 4) and the sexual Chunchon and 180 clones at Yongin with a space of 4mx4m. asymmetry (Eq. 5) were estimated using Choi and others’ Reproductive characteristics of 161 clones, which the two (1999) methods as follows: clone banks have in common, were investigated for 3 con- n secutive years (1998 to approximately 2000). ¤ 2 –1 m = ( xi() ) (1) The clone banks were not considered as fully mature åi=1 populations when the numbers of female and male strobili were counted. Generally, Korean pine begins to show strobili at age of 12 or 15 in natural stands. Grafted clones, however, n ¤ 2 –1 produce strobili earlier than natural stands. In these clone m = { å xi( ) } (2) banks, there is not much difference in height (4 to 5m) and i=1 DBH. Five ramets per clone were chosen for assessment in 1 early June. The number of female strobili was counted ma = ⁄2 (m + m ) (3) individually from a whole tree. The total number of male strobili was estimated by multiplying the average number of n 1 2 –1 strobili per branch by the total number of branch bearing mb = { ( ⁄2(xi()+xi( )) } (4) male strobili. åi=1 Analysis of variance (ANOVA) tests and heritability esti- mates were conducted based on the data for female and male As = ma /|mb – ma|, 0 £ As £ 1 (5) strobili production. The ANOVA was performed using a logarithmic transformation of the original data to normalize where n is the total number of clones, m is the female the distribution of variances (Steel and Torrie 1980). SAS effective population number, and xi () is the proportion of program (ver 6.12; SAS Institute Inc., 1996) was used for the female strobili of the ith clone to the whole production of ANOVA tests and heritability estimation. Broad-sense heri- females. m and xi ( ) in males correspond to those for 2 tabilities (H ) were estimated on the basis of individual trees females. ma is the arithmetic mean of the two measures (m (Schmidtling 1983) as: and m ) and mb is based on the relative frequency of xi () and xi . In this study, we used the relative effective population 2 ( ) H 2 s c . number instead of effective population number for easy = 22 comparison with those of other studies. ssce+ Parental balance was assessed using a cumulative gamete contribution curve (Griffin 1982). The numbers of female Results and Discussion __________ and male strobili were ordered by clone from high to low strobilus production, and the cumulative contribution per- Reproductive Characteristics and centages were plotted against the proportion of the clones Heritability (Kang 2000). The maleness index (Ai) is defined as the proportion of a Large variations in both female and male strobilus pro- clone’s reproductive success that is transmitted through its duction among clones were observed at both Yongin and pollen (Kang 2000). Maleness index based on strobilus Chunchon (table 1). The differences of male strobilus pro- production was estimated as follows: duction among clones were far more extreme than that of ai female strobilus production. It seems that this phenomenon Ai = is a typical character of Korean pine from our experience of gi + ai Table 1—Mean, standard deviation (S.D.) and coefficient variation (C.V.) for the number of female and male strobili at Yongin and Chunchon during the period of 1998 to 2000. Yongin Chunchon 1998 1999 2000 1998 1999 2000 Female Male Female Male Female Male Female Male Female Male Female Male Mean 5.3 64 13.6 240 9.1 36 2.2 146 12.5 393 5.8 307 S.D.(±)4.6184 12.8 975 9.2 111 2.1 514 9.3 988 5.1 982 C.V.( percent) 87 289 94 406 101 305 97 353 75 251 87 320 USDA Forest Service Proceedings RMRS-P-32.
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