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Growth and Phenology of Hybrid Aspen Clones (Populus Tremula L

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Growth and Phenology of Hybrid Aspen Clones (Populus Tremula L Silva Fennica 35(1) research articles Growth and Phenology of Hybrid Aspen Clones (Populus tremula L. × Populus tremuloides Michx.) Qibin Yu, P. M. A. Tigerstedt and Matti Haapanen Yu, Q., Tigerstedt, P.M.A. & Haapanen, M. 2001. Growth and phenology of hybrid aspen clones (Populus tremula L. × Populus tremuloides Michx.). Silva Fennica 35(1): 15–25. Height, basal diameter, diameter at breast height, bud burst, and leaf development were recorded in a 5-year-old hybrid aspen clonal trial. The fi eld trial consisted of four aspen hybrid clones (Populus tremula × Populus tremuloides ) and one local P. tremula seedling source. Phenological traits were observed in the 3rd year. Growth patterns were recorded during the 3rd and 4th years. Phenological traits were explored in relation to hybrid vigor expressed as growth traits. Differences were observed for phenological and growth traits among hybrid clones and P. tremula. The growth period varied from 143–158 days for the four hybrid clones, and was 112 days for P. tremula. The correlation between growth period and yield was highly signifi cant. The annual growth rate of height for the hybrids was 4.2 cm per 7 days (2.4 for P. tremula) in the 3rd year and 6.4 cm per 7 days (2.9 for P. tremula) in the 4th year. After 5 years, mean estimated stem volume of the hybrids was 3.9 times that of P. tremula. Signifi cant clone by year interaction was observed for height, diameter, and volume growth. The hybrid vigor seems to be mainly attributable to a longer growth period. Keywords hybrid aspen, growth pattern, growth rate, phenology Authors’ addresses Yu and Tigerstedt, Department of Plant Biology, P.O. Box 27, FIN-00014 University of Helsinki, Finland; Haapanen, Finnish Forest Research Institute, P.O. Box 18, FIN-01301 Vantaa, Finland E-mail qibin.yu@helsinki.fi Fax +358 9 7085 8727 Received 24 March 2000 Accepted 8 January 2001 1 Introduction in high-quality papermaking. The Finnish forest industry has shown a fresh interest in utilising Populus tremula is a widely distributed species aspen for producing the short fi bres. Interest in in Finland, often growing in mixed stands with cultivating selected clones of hybrid aspen has pine, spruce and birch. Recent developments in increased. The hybrid between European aspen P. wood fi bre research have allowed short fi bres to tremula L. and North American trembling aspen be used in admixture with coniferous long fi bres P. tremuloides Michx. has shown superior growth 15 Silva Fennica 35(1) research articles in Finland (Beuker 1989). Genetic gain from Breeding and the University of Helsinki. The trial aspen breeding programs were demonstrated in is located in Viikki, area of Helsinki University the United States (Einspahr 1984) and Europe (lat. 60°14´ N, long. 25°05´ E, alt 10 m). The (Melchior 1985). Interspecifi c hybrids have trial was planted in May, 1994 with one-year-old grown faster than the progenies of intraspecifi c plants, and consists of four aspen hybrid clones crosses (Heimburger 1936, Einspahr and Benson (Populus tremula × P. tremuloides) and one P. 1964, Melchior and Seitz 1966, Heimburger tremula seedling source. Clone 1 was a hybrid 1968, Zsuffa 1969, 1973). There is much genetic between P. tremula (Tuusula, south Finland) × P. diversity in aspen, and it can be manipulated tremuloides (Lake Aleza, B.C. Canada). Clone 2 through selective breeding, interspecifi c hybridi- was a hybrid between P. tremula (Tuusula, Fin- zation and cloning (Li 1995). land) × P. tremuloides (Maple, Ontario, Canada). Hybrid vigor has been observed in poplar and Clones 3 and 4 were selected from a mixed plan- has been one of the driving forces in poplar breed- tation of hybrids, between P. tremula (Tuusula, ing (Muhle Larsen 1970). The phenomenon is also or Punkaharju, Finland) and P. tremuloides (Lake well documented for aspen hybridisation in the Aleza or Maple, Canada). Thus their exact ped- U.S. (Li and Wu 1996, 1997, Li et al. 1998). This igrees are unknown. The local aspen seedlings increase in vigor may have many components, were progenies of single open pollinated superior including carbon allocation patterns, water and tree in Karkkila (lat. 60°32´ N, long. 24°15´ E), nutrient use effi ciency, and shoot growth phenol- Finland. Thus they are predominantly half-sib. A ogy. Traits that affect Populus performance include randomized complete block design was used with phenology (Michael et al. 1990, Ceulemans 1992), 4 × 4 plants spaced at 2.5 m × 2.25 m per plot and leaf and stomatal morphology and photosynthetic fi ve blocks. capacity. Large clonal differences in phenology, Total height, basal diameter 15 cm from the leaf area development, and photosynthetic produc- ground and breast height diameter at height 1.3 tion have been found (Ceulemans 1987, Orlovic m were measured. Measuring positions were 1998, Thomas et al. 1997). Li et al. (1998) found marked on the stems to assure that repeated meas- that interspecifi c aspen hybrids grew faster than urements were taken at exactly the same position. intraspecifi c hybrids at the juvenile stage. The Stem volume was estimated as height × basal authors ascribe this to larger internode number and diameter2 (Causton 1985, Ceulemans 1992, Li length and leaf number. Heterosis of P. tremula × et al. 1998). P. tremuloides hybrids for stem volume was prob- Height, basal diameter and breast height diam- ably the result of delayed bud set, which gave a eter were measured in the autumns of the 2nd to longer duration of height growth. the 5th year. For observation of dynamic seasonal In this paper we compare hybrid aspen clones growth patterns, the fi ve tallest trees were sampled and local P. tremula for yield, growth pattern and for measurements of growth traits in each plot in phenology. The aim of the study was to obtain the spring of the 3rd year. Height, basal diameter better fundamental understanding of hybrid vigor and breast height diameter were measured at of aspen hybrids. We present results of studies on intervals of approx. 2 to 3 weeks over the course of yield components and their relationship in hybrid the growing season in the 3rd year (10 measure- aspen clones and local P. tremula. The growth ments) and the 4th year (8 measurements). The patterns were studied mainly from a phenological growth rates for height, basal diameter and breast perspective, in an attempt what has hitherto been diameter were expressed as increments (cm or loosely termed heterosis of species hybrids. mm) over a 7 day period. The annual growth rates were calculated by bringing total growth made over the total growth measuring period (days) to a 7 day basis. 2 Materials and Methods Phenological traits were recorded in the 3rd year. Growth initiation was classifi ed into three The material used in this study was a clonal trial stages counting from the fi rst of May: 1) the day established by the Foundation for Forest Tree when the bud emerged 5 mm beyond the bud 16 Yu, Tigerstedt and Haapanen Growth and Phenology of Hybrid Aspen Clones... scale, 2) the day leaves unfolded, 3) the day the is the interaction of the i th clone and k th block leaves attained full size. Likewise, three stages of in the j th year and εijkl is random error. Clonal growth cessation were distinguished: 1) the day effects were considered to be fi xed, while the year when half of the leaves yellowed, 2) when all of and block were treated as random effects. the leaves yellowed, 3) when all leaves defoliated. “Bud emerged” and “half of the leaves yellowed” were taken as indicators of growth initiation and growth cessation. The length of the growth period 3 Results was estimated as the number of days from growth initiation to growth cessation. Each individual 3.1 Growth was observed from the same direction and from about the same relative position with respect to Highly signifi cant differences in growth traits the tree crown, every other day in the spring, and (height, basal and stem volume) were observed every third day in the autumn. among the blocks, clones, years and their inter- All calculations were based on plot means. actions (Table 1). All hybrid clones had out- The difference between the hybrid clones and performed the P. tremula by the end of the 5th the P. tremula were established by an analysis year. The best clones were clone 2 and 1, with of variance (ANOVA) using the PROC GLM stem production volumes of 17.82 dm3 and 17.57 procedure of the SAS statistical software pack- dm3, respectively (Fig. 1). Clonal ranking in the age (1989; SAS Institute, Cary, NC) with the 5th year remained nearly the same as in the 2nd type III estimation of sum of squares. Means of year. Clone 3 was shorter than clone 4 at the end phenological and growth traits were separated of the 5th year, but because of its larger diameter, by Tukey’s multiple range test at p ≤ 0.05 prob- it had a larger stem volume (Table 2). ability level of signifi cance. Pearson correlations Growth patterns of mean height, basal and were calculated to assess the linear relationships breast diameter of the hybrid clones and P. trem- between the studied traits. Data from all the four ula in the 3rd and the 4th year are shown in Fig. years were combined and the following statistical 2. The hybrids displayed superior growth for all model was used: traits during the entire growth period (Table 3). The growth rate of height, basal and breast Xijkl = µ + Ci + Yj + Bk(j) + CiYj + CiBk(j) + εijkl height diameter was highest from the beginning of June to the end of July in the 3rd year.
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