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Juvenile - mature wood transition in second-growth coastal Douglas-fir Di Lucca, Carlos Mario 1987

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JUVENILE - MATURE WOOD TRANSITION IN SECOND-GROWTH COASTAL DOUGLAS-FIR By CARLOS MARIO DI LUCCA Eng., Universidad Nacional de La Plata, 1978 THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF THE FACULTY OF GRADUATE STUDIES Department of Forestry We accept this thesis as conforming to |the required standard THE UNIVERSITY OF BRITISH COLUMBIA July 1987 ccjCarlos Mario Di Lucca, 1987 MASTER OF SCIENCE i n In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date AUGUST 7 / 3?1 DE-6(3/81) ABSTRACT The transition from old-growth to second-growth British Columbia coastal Douglas-fir has resulted in reduction of log size and increased proportion of juvenile (core or crown-formed) wood. Determination of the zone of transition from juvenile to mature wood is critical to the definition of wood quality and timber value. Thirteen unpruned, two pruned second-growth, and two unpruned plantation-grown coastal Douglas-fir trees were sampled to analyze the hypothesis that the transition in relative density from juvenile to mature wood occurs at the base of the live crown. X-ray densftometric techniques were utilized to determine yearly pith to bark relative density data of five cross-sectional discs from each tree. Segmented linear regression techniques were utilized to estimate the juvenile - mature wood transition age from the data. The average number of growth increments from the pith at which juvenile - mature wood transition occurred on sections sampled at breast height, 20 percent and 40 percent of total height was 22.18. When the hypothesis was tested on unpruned trees, before and after harvest, the juvenile - mature wood transition occurred below the base of the live crown. When the hypothesis was tested on pruned trees, the transition occurred at the base of the live crown, which represented the upper limit of pruning height. This information may provide a greater insight into juvenile - mature wood transition. It wi11 likely assist in the determination of wood quality and economic value of forest products manufactured from second-growth coastal Douglas-fi r. f V TABLE OF CONTENTS Page TITLE PAGE 1 ABSTRACT i i TABLE OF CONTENTS iv LIST OF TABLES vi LIST OF FIGURES viiLIST OF APPENDICES xACKNOWLEDGEMENT i i i 1.0 INTRODUCTION 1 2.0 LITERATURE REVIEW 5 2. 1 Wood Format i on2.1.1 Primary Growth 5 2.1.2 Secondary Growth 7 2.1.3 Influence of the Crown on Wood Formation .. 10 2.2 Juvenile and Mature Wood 13 2.2.1 Anatomical Structure Comparison 15 2.2.2 Physical Properties Comparison 17 2.2.2.1 Relative Density 19 2.2.2.2 Other Properties 24 2.2.3 Juvenile - Mature Wood Transition Determination 26 2.2.3.1 Subjective Methods 27 2.2.3.2 Objective Methods 29 V 3.0 MATERIALS AND METHODS 33 3.1 External Tree Characteristics 34 3.2 Internal Tree Characteristics 6 3.2.1 X-ray Dens i tometr i c Analyses 37 3.2.2 Juvenile - Mature Wood Transition Determination 38 4.0 RESULTS AND DISCUSSION 43 4.1 X-Ray Densitometric Analyses 44.2 Juvenile - Mature Wood Transition Determination .. 46 4.2.1 Juvenile - Mature Wood Relative Density Values 50 4 . 3 Base of the L i ve Crown 51 4.4 Relationships Between Juvenile - Mature Wood Transition and the Base of the Live Crown 54 4.4.1 Hypothesis Testing Before Harvest 5 4.4.2 Hypothesis Testing After Harvest 63 4.5 Pruned Trees 68 5.0 CONCLUSIONS 72 LITERATURE CITED 7 APPENDICES 130 vi LIST OF TABLES Table Page 1 Douglas-fir stand characteristics 91 2 Sample tree characteristics 92 3 Tree crown characteristics 3 4 Summary of relative density values for sections sampled at breast height (B.H.) 94 5 Summary of relative density values for sections sampled at 20 percent of total height 95 6 Summary of relative density values for sections sampled at 40 percent of total height 96 7 Summary of relative density values for sections sampled at 60 percent of total height 97 8 Summary of relative density values for sections sampled at 80 percent of total height 98 9 Summary of the distribution of relative density profiles which showed no Juvenile - mature wood transition 99 10 Transition determination by comparing simple against segmented linear regression models on relative density profiles 100 11 Summary of the distribution of relative density profiles which showed juvenile - mature wood transition 102 12 Summary of the relative density values of the profiles which showed no juvenile - mature wood transition 103 13 Summary of the relative density values of the profiles which showed juvenile - mature Wood transition 104 14 Overall summary of relative density values for juvenile and mature wood 105 vi i 15 Summary of sampled dead branch characteristics .... 106 16 Summary of lowest relative density values 107 17 Differences between height of juvenile - mature wood transition point, height of crown base and total height 108 18 Summary of height difference statistics 109 19 Correlation matrix for live crown base height, average crown height, average transition height A and average transition height B 110 viii LIST OF FIGURES F igure Page 1 Location of Douglas-fir stands Ill 2 Pith to bark relative density profiles for sample tree 1A5 112 3 a) Segmented linear regression model on pith to bark relative density profile for a sample radi us 113 b) Simple linear regression model on pith to bark relative density profile for a sample radius4 a) Scatter plot of height over number of growth increments at which juvenile - mature wood transition occurs 114 b) Scatter plot of height over number of growth increments at the base of dead branches for all trees 11c) Scatter plot of height over number of growth increments which indicates the lowest relative density value in the profile 114 5 a) Scatter plot and height prediction model for crown base height over total height 115 b) Scatter plot and height prediction model for average crown height over total height 115 6 Scatter plot of a) total height and height prediction models over total height; for: b) average crown height; and c) crown base height ... 116 7 Graphical representation of: a) total tree height; and b) crown base height over number of growth increments from the pith 117 8 Graphical representation of: a) total tree height; b) crown base height before harvest; c) live crown base height at harvest; and d) average crown height at harvest over number of growth increments from the pith at breast height 118 fx 9 a) Scatter and height prediction model for total tree height over number of growth increments from the pith at breast height 119 b) Scatter and height prediction model for lowest relative density over number of growth increments from the pith at breast height 119 10 a) Scatter and height prediction model for height to crown base over number of growth increments from the pith at breast height 120 b) Scatter and height prediction model for juvenile - mature wood transition points over number of growth increments from the pith at breast height 120 11 Height prediction models for: a) total tree height; b) lowest relative density height; c) crown base height; d) Juvenile - mature wood transition height over number of growth increments from the pith at breast height; and e) diagramatic tree representations 121 12 a) Scatter and height prediction model for the difference between crown base height and juvenile - mature wood transition height over juvenile - mature wood transition height . 122 b) Scatter and height prediction model for the difference between total height and juvenile - mature wood transition height over juvenile - mature wood transition height . 122 13 Graphical representation of: a) estimation of the average transition height A; and b) estimation of the average transition height height B over number of growth increments from the pith at breast height 123 14 Scatter plot of juvenile - mature (J. M.) wood transition number of growth increments over tree sections and overall section averages, i.e. average transition age A 124 15 Scatter plot of height differences among total and transition heights over tree sections and overall section averages, i.e. average transition height B 125 16 a) Scatter plot and height prediction model for average transition height A over total height . 126 b) Scatter plot and height prediction model for average transition height B over total height . 126 X 17 Scatter plot of a) total tree height, and height prediction models over total height for: b) average crown height; c) average transition height A; d) live crown base height; and e) average transition height B 127 18 Pith to bark relative density profiles for pruned tree RF2 128 19 Pith to bark relative density profiles for pruned tree RF3 129 xi LIST OF APPENDICES Appendi x Page 1 Average Crown Height. Estimation Procedure 130 a) Relationship between total branch length (BL) and the vertical distance from the leader (L) for Sample Tree 1A5 132 b) Relationship between total branch length (BL) and the transformed vertical distance from the leader In [ (L/c)+l ] for Sample Tree 1A5 . 133 c) Average crown height position determination for Sample Tree 1A5 134 2 Relationship between total height and branch age to estimate base of live crown positions at young ages 135 3 Summary of X-Ray Dens?tometric Analysis Procedure 136 4 Fortran program to determine residual sum of squares using non-linear optimization routines for segmented regression models 138 5 Tree 1A5 139 6 Tree 1A7 140 7 Tree 1B6 1 8 Tree 1B11 142 9 Tree 1C1 3 10 Tree 1C6 144 11 Tree 1D5 5 12 Tree 1D6 146 13 Tree 1E3 7 14 Tree 1E8 148 15 Tree 1F7 9 xi i 16 Tree IF 10 150 17 Tree CR 1 5 1 18 Tree CR2 152 19 Tree RF 1 3 20 Summary of relative density data: a) Tree 1A5 154 b) Tree 1A7c) Tree 1B6 5 d) Tree IBM 15e) Tree 1C1 6 f) Tree 1C6g) Tree 1D5 157 h) Tree 1D6i) Tree 1E3 8 j) Tree 1E8 15k) Tree 1F7 9 1) Tree IF 10m) Tree CR1 160 n) Tree CR2o) Tree RF 1 1 p) Tree RF2 16q) Tree RF3 2 x i i i ACKNOWLEDGEMENT The author acknowledges, with gratitude, Dr. J.W. Wilson, Faculty of Forestry, The University of British Columbia, Dr. R.M. Kellogg, Forintek Canada Corporation and Dr. K.J. Mitchell, Ministry of Forests and Lands of British Columbia, under whose guidance and supervision this project was accomplished. Appreciation is also due to Dr. P. Marshall, Faculty of Forestry, The University of British Columbia, for his advice and review of the thesis. Grateful acknowledgement is given to L.A. Jozsa, S.G. Johnson, J. Cook and J. Richards, Forintek Canada Corporation, for their valuable assistance during experimental phases of the study. Appreciation is extended to Forintek Canada Corporation for the use of their laboratory facilities and materials. Special thanks is due to J.C. Fahler, Est. Las Marias, Argentina, for inspiring the author's interest in the study of forestry. The deepest gratitude is expressed to my wife, Barbara, for her patience and encouragement. Finally, the author is appreciative of the financial support provided by the Research Council of British Columbia, the Ministry of Forests and Lands of British Columbia and the Canadian Forest Service. 1 1.0 INTRODUCTION Coastal Douglas-fir (Pseudotsuga menz i es i i (Mirb.) Franco) is found west of the Cascade Range in Washington and Oregon, west of the Coast Range in British Columbia and west of the Sierra Nevada in northern California. The north - south range of the species extends approximately 3400 km south from the central British Columbia coast (Lat. 55°N) (Fowells, 1965). Coastal Douglas-fir is a relatively minor species in British Columbia. Reserves of coastal Douglas-fir in the Vancouver and Prince Rupert regions amount to approximately 126 million cubic metres, which represent six percent of the total mature timber and six percent of the annual timber harvested in 1980 (British Columbia Ministry of Forests, 1980). However, because of easy access, high quality and developed markets, Douglas-fir is highly demanded and the most intensively managed commercial species. For these reasons, the old-growth coastal Douglas-fir resource has been almost replaced by young and fast grown second-growth timber. The transition to second-growth timber has resulted in reduction of log size and changes in the quality of the raw material. These changes are demonstrated by the increased proportion of juvenile (core or crown-formed) wood found in 2 the second-growth timber. Compared to mature (stem-formed) wood, the juvenile wood of Douglas-fir is characterized by lower relative density, shorter tracheids, larger fibril angle, lower strength, lower percentage of latewood, lower transverse shrinkage, higher longitudinal shrinkage, lower cellulose content and higher lignin content (Bendtsen, 1978; Barrett and Kellogg, 1984, 1986; Jackson and Megraw, 1986; Jozsa and Kellogg, 1986; McKimmy, 1986). Such differences in wood properties are very important in the processing and manufacturing of lumber, veneer, chips and other forest products. The future of the Canadian forest industry will depend upon the successful conversion and commercialization of forest products manufactured from second-growth timber. In order to maintain and improve wood supply the industry needs information to plan and predict the effects of intensive forest management on volume production, properties and value of second-growth timber (Kellogg, 1986). The Tree and Stand Simulator (TASS) (Mitchell, 1975; Mitchell and Cameron, 1985) models the growth and yield of second-growth coastal Douglas-fir under different management conditions. This is a biologically oriented model which can simulate crowns of individual trees in a three-dimensional growing space in response to internal 3 growth processes, environmental factors, physical restrictions and cultural practices. However, the model does not consider wood qualities or value of wood products. In view of the above, Forintek Canada Corporation, the Pulp and Paper Research Institute of Canada, the Ministry of Forests and Lands of British Columbia and the University of British Columbia have initiated nine integrated projects to provide much of the technical information required for future utilization planning for second-growth coastal Douglas-fir (Kellogg, 1986). The purpose of this thesis project, which is one of the nine Douglas-fir Task Force projects, fs to investigate the transition in relative density between juvenile and mature wood. The specific objective of this study is to test and analyze the hypothes i s that: The relative density transition from Juvenile to mature wood in individual growth increments of second-growth coastal Douglas-fir trees occurs at the base of the live crown. To test this hypothesis, 13 unpruned, two pruned second-growth, and two unpruned plantation-grown trees were sampled and analyzed. 4 It fs hoped that the results of this project will contribute to the understanding of how and when the relative density transition between juvenile and mature wood occurs. Furthermore, the results can help extend TASS to simulate the production of juvenile and mature wood, thus providing insight into interactions of stand dynamics and wood quality. In addition, the findings of this project are likely to be of value to the Douglas-fir Task Force and the general forestry industry in the determination of wood quality and, ultimately, economic value of forest products in British Columbia. 5 2.0 LITERATURE REVIEW 2.1 Wood Formation The physiological processes controlling wood formation in temperate zone trees are related to the seasonal activity of the vascular cambium and differentiation of its der i vat i ves. In coniferous trees, there is a high correlation between growth and development of the crown over time and wood formation along the stem (Larson, 1964, 1969). The physiology of wood formation must, then, be considered as an integral part of tree growth. This growth can be divided into two developmental stages, a primary stage originating in the vegetative terminal buds, and a secondary stage originating in the vascular cambium. 2.1.1 Primary Growth The primary growth stage encompasses the elongation of the main stem and branches and the regulation of height growth and tree form. In temperate zone conifers, the seasonal primary growth begins as a result of an increase in temperature and day length in early spring (Wilcox, 1962; Panshin and de Zeeuw, 1980). These changes in 6 environmental conditions initiate primary cell division after the rehydration, swelling and increase of hormonal and enzymatic activity in the apical men"stems inside the vegetative buds. In Douglas-fir, this growth activation occurs in the last week of March at lower elevations (Owens, 1968; Allen and Owens, 1972). As the growing season advances, more new cells are formed causing primary shoot elongation in the stem and branches. Subsequently, the newly formed cells undergo differentiation, resulting in changes in size, shape and function. In this manner, primary permanent tissues are differentiated from three meristematic tissues called protoderm, ground meristem and procambium (Kowlowski, 1971). The protoderm develops an external protective layer called the epidermis. The ground meristem develops into the central pith and cortex. The procambium gives rise to the primary phloem and primary xylem and subsequently the vascular cambium. The primary phloem is formed on the outer side of the procambium. It performs the function of mobilization and transport of sugar and other nutrients within the shoot apex. The primary xylem is formed on the inner side of the procambium and it performs the function of support and conduction of dissolved substances from the roots to the terminal shoots. 7 The vascular cambium, a secondary meristematic tissue responsible for the lateral growth of xylem (wood), and phloem (bark) tissues, is formed after completion of the differentiation of primary permanent tissues. It is made up of fusiform and ray cambial initial cells, and separates the xylem and phloem mother cells. 2.1.2 Secondary Growth The secondary growth stage, as it relates to wood formation, can be summarized into the following developmental phases (Larson, 1969; Brown, 1970; Kozlowski, 1971; Wilson, 1984; Haygreen and Bowyer, 1982): 1. Awakening, rehydration and swelling of the dormant cambium at the beginning of the growing season; 2. Periclinal division of fusiform cambial initial cells resulting in the formation of new meristematic cells, and new xylem and phloem mother cells with cell division peaking a few weeks into the growing season, after which it decreases gradually until completed in early summer; 8 3. Differentiation of the xylem and phloem cambial mother cells passing through the phases of enlargement, and secondary wall formation and thickening; and 4. Finally, maturation of the newly formed xylem and phloem cells, thus completing the secondary wall thickening and 1ignification phases. The annual addition of newly formed secondary phloem cells forms the bark, which is divided into a light coloured inner living bark and a dark outer dead bark. The annual additions of secondary xylem cells form concentric layers or rings of growth increments that are responsible for the increase in stem and branch diameters (Panshin and de Zeeuw, 1980). The annual growth increments are characterized by the formation of two zones called ear 1ywood (spr i ngwood) and 1 atewood (summerwood). The most important anatomical differences between ear 1ywood and latewood are the radial cell diameter and the secondary wall thickness. Such characteristics are known to be independent of each other (Richardson and Dinwoodie, 1960; Wodzicki, 1960). Their formation, particularly in pine trees, was explained by Wareing (1958) and Larson (1963, 1969) in the following two hypotheses. 9 The first is the hormonal or auxin hypothesis, which is related to the regulation of tracheid diameter. During the period of active shoot elongation and needle development in the spring, a high level of diffusible auxin is produced within the vegetative buds and transported downward along each branch, down the main stem and into the roots. Then, the cambial activity begins with the production of large diameter xylem tracheids classified as earlywood. Earlywood formation continues as long as the elongating shoots and developing needles compete for the stored and currently produced photosynthates. As the growing season advances, narrow diameter latewood tracheids are produced in response to the cessation of terminal growth, the reduction of auxin synthesis, and the increase of growth inhibiting substances. The formation of latewood tracheids begins at or near the base of the tree and tapers upwards to a point of extinction near the apex. The second hypothesis explains the secondary cell wall thickening, which is related to the net amount of photosynthates that reach each tracheid after respiration requirements have been met. The competition for photosynthates among meristematic tissues decreases, and the rate of photosynthesis in the current year needles rapidly increases (Freeland, 1952; Clark, 1961). Thus, more photosynthates are available for both stem growth and 10 secondary wal1 thickening of the latewood tracheids (Rutter, 1957). 2.1.3 Influence of the Crown on Wood Formation The hormonal and the secondary wal1 thickness regulation hypotheses emphasize that earlywood and latewood tracheid formations are correlated to photosynthate availability and presence of auxin. These crown formed products determine the rate of cambial initial cell division and the degree of differentiation of the cambial derivatives. It fs clear that the growth and development of the crown over time has a direct regulatory influence over wood formation in the stem (Larson, 1962, 1964). Within the crown, the most vigorously active live branches will regulate the wood formation by a steady production of photosynthates and auxins. These branches are characterized by having complete growth increments along their length forming an active union with the main stem. As the branches start to compete for light and become older and longer, their vigor, given by the capacity of producing photosynthates and auxins, gradually decreases (Larson, 1969). This leads to a decrease in the current shoot length from the apex to the base of the live crown and from the tip to the base of the branch (Fraser, 1962; Forward and Nolan, 1964). 11 The decrease in branch vigor is followed by a period of senescence which starts at the lower crown branches. At this time the contribution of these branches to wood formation gradually decreases, leading to the formation of incomplete or absent growth increments at the branch base (Andrew and Gill, 1939; Reukema, 1959, 1961; Reeb, 1984). These senescent branches, which are still attached to the main stem, interrupt their contact with the main translocation pathways in the tree, resulting in diminishing contribution to wood formation until branch death occurs. Larson (1969) concluded that major changes in wood formation and quality occur below the living crown, or below the most active branches of the crown. For tree growth analysis purposes, the height to live crown base has been defined in several ways. Smith et a 1. (1961) and Mitchell (1969, 1975) defined the height to live crown as the average distance from the ground to the lowest live branch in each of four quadrants. They also defined an average live crown height as the average height from the ground to the point of maximum crown spread or crown radius. This point occasionally represents the crown contact with the crowns of neighbouring trees. Reeb (1984) defined height to live crown base as the height from the ground to the whorl at which at least 75 percent of the branches are alive. 12 The number of growth increments at the base of the dead branches and the corresponding height positions can be utilized to determine the approximate crown base positions at younger ages. The age of the tree also has an influence on wood formation. A young tree with a high percentage of the stem covered with active live branches will produce wide growth increments with a high proportion of earlywood trachelds. This is explained by the prolonged influence of the apical meristems on the cambial regions in the active live crown. As the tree grows older and distance from both the pith and the active live crown increases, the proportion of latewood tracheids in the growth increments gradually increases. Now, the cambial regions below the crown become less influenced by the apical meristems. Consequently, two characteristic wood zones can be differentiated within the tree stem as a function of distance from the active live branches and number of growth increments from the pith. The first zone, called juvenile wood, forms a central core of wood around the pith extending from the base to the apex of the tree. The second, called mature wood, is formed around the juvenile wood core and below the living crown. 13 2.2 Juvenile and Mature Wood A universal definition of the terms juvenile and mature wood has not been agreed upon, although such terms can arbitrarily describe the type of wood produced in relation to crown proximity and number of growth increments from the pith. Several authors have defined juvenile and mature wood from different points of view. For instance, when considering the juvenile and mature wood stem positions in a tree, juvenile wood has been called core, inner, or pith-associated wood and mature wood has been called exterior, outer or non-pith-associated wood (Perry and Wang, 1958; Zobel et a]_. , 1959; Moody, 1970). Juvenile wood has been called immature or youthful wood and mature wood has been called old or adult wood, based on their respective physiological stages of maturity (Rendle, 1958, 1959, 1960). Juvenile wood has been called crown-formed wood, because it is formed inside the living crown, and mature wood has been called stem-formed wood due to its formation outside of the living crown (Trendelenburg, 1935; Cooper, 1960; Brunden, 1964; Larson, 1969, 1973). The size of the juvenile wood core generally depends upon the growth rate, regardless of the species. Si 1vicultural practices such as fertilization, irrigation 14 and thinning will tend to decrease crown recession and increase crown vigor, growth rate and the size of the juvenile wood core (Bendtsen, 1978; Briggs and Smith, 1986; Megraw, 1985; Oliver, 1986). The proportion of the juvenile wood core in the stem is a function of the species, number of growth increments from the pith and distance from the active live crown (Paul, 1960; Bendtsen, 1978). This proportion tends to be high in early harvested trees and in open-grown trees. Increasing crown recession by delaying the harvesting age of stand-grown trees will decrease the juvenile wood proportion at harvest age. An abrupt change from juvenile to mature wood can also be achieved by pruning the live and vigorous lower branches in the crown (Marts, 1949; Gerischer and De Vilifers, 1963; Smith, 1968; Larson, 1965, 1969; Cown, 1973; Polge et aj_. , 1973; Plumptre and Austin, 1978). It is evident that growth and development of the active live crown as a function of age will determine not only the characteristics of the tracheids formed, but also the quantity and quality of juvenile and mature wood in the stem. Juvenile and mature wood must be considered as two different populations within the same tree because of their fundamental differences in wood quality (Panshin and de 15 Zeeuw, 1980). Such differences are determined primarily by the tracheids' anatomical structure and chemical composition, and secondarily by the derived physical wood propert i es. 2.2.1 Anatomical Structure Comparison When considering the tracheid anatomical structure variations in stem cross-sections of most conifers, the length, diameter and secondary wall thickness of the juvenile wood tracheids increase progressively from the pith until they more or less stabilize in the mature wood (Bendtsen, 1978; Panshin and de Zeeuw, 1980; Megraw, 1985; Krahmer, 1986). In softwoods, the length of mature wood tracheids can be up to three or four times greater than that of juvenile wood tracheids (Anderson, 1951; Dadswel1, 1958; Dinwoodie, 1961). Many studies have shown that the tracheids in juvenile wood are shorter than in mature wood, regardless of height. These results were reported by Zobel and Kellison (1972) for loblolly pine (Pinus taeda L.), by Loo et al. (1985) for slash pine (Pinus e11iotti i Engelm.), by Wang and Micko (1984) for white spruce (Picea g1auca (Moench.) Voss), by Erickson and Harrison (1974) and Jackson and Megraw (1986) for Douglas-fir, by Boone and 16 Chudnoff (1972) for Caribbean pine (Pinus caribaea Morelet.), by Cown (1975) for radiata pine (Pinus radiata D.Don), and by Well wood and Jurazs (1968) for western redcedar (Thu.ia p 1 icata Donn.). The secondary wall thickness of the juvenile wood tracheids was reported to be less than that of mature wood tracheids by Zobel and Kellison (1972) for loblolly pine, by Isebrands et al_. (1982) for larch (Larix spp.), by Cown (1975) for radiata pine and by Foelkel et al.. (1976) for slash pine. Erickson and Harrison (1974) found that in Douglas-fir the radial and tangential diameter of the juvenile wood tracheid increased toward mature wood. In addition to the above findings, the lumen size, and the microfibril angle of the S2 layer of the juvenile wood tracheids generally decrease toward mature wood, where they stabilize to some extent. The decrease in lumen size from juvenile to mature wood tracheids was reported by Zobel and Kellison (1972) for loblolly pine, by Foelkel et aj.. (1976) for slash pine, and by Cown (1975) for radiata pine. The decrease in microfibril angle from juvenile to mature wood tracheids was found by Meylan (1968) for radiata pine, by Boone and Chudnoff (1972) for Caribbean pine, and by Erickson and Arfma (1974) for Douglas-fir. 17 Variations in the chemical composition of tracheids include the following. The holocellulose and alpha cellulose contents of the juvenile wood tracheids gradually increase from the pith until they begin to stabilize in the mature wood. However, an inverse relationship occurs with some hemicelluloses and lignin, which decrease from juvenile to mature wood. These trends can be seen from the data of Kirk et aj_. (1972) and Zobel and Kellfson (1972) on loblolly pine, Schmidt and Smith (1961) on Caribbean pine. Well wood and Smith (1962), Kennedy and Jaworsky (1960), Sastry and Wei 1 wood (1971), Erickson and Harrison (1974), and Megraw (1985) on Douglas-fir. In conclusion, in comparison to mature wood, the juvenile wood in Douglas-fir is characterized by smaller and shorter tracheids with thinner walls and larger microfibril angles, and by lower holocel1ulose and alpha cellulose contents and higher lignin and hemicellulose contents. 2.2.2 Physical Properties Comparison Anatomical and chemical differences among tracheids, as described, influence the physical properties and, therefore, the quality of juvenile and mature wood. To define wood quality, Larson (1969) stated: 18 "During the wood formation process numerous factors both inside and outside the tree lead to variation in type, number, size, shape, physical structure and chemical composition of the wood elements. Wood quality is an arbitrary classification of these variations in the wood elements when they are counted, measured, weighed, analyzed or evaluated for some specific purpose." The quality of wood, then, refers to its fitness for particular utilization, each quality being determined by a certain number of properties. For instance, relative density, strength, elasticity, proportion of corewood, reaction wood, grain orientation, permeability, moisture content and presence of knots are some of the properties which determine the suitability of wood for a specific end use (Fielding, 1967; Haygreen and Bowyer, 1982). When considering softwood as a raw material source fo pulp and lumber production, relative density is one of the most important characteristics to be considered as a general indicator of wood quality. The reason is that relative density is highly correlated to pulp yield, paper-making properties and strength properties of timber (Barefoot et aj_. , 1970; Gonzalez and Kellogg, 1978; Kellogg, 1982;). 19 2.2.2.1 Relative Density Relative density, often called specific gravity in wood quality studies, expresses how much cell wall substance is present in a given volume of wood. Relative density is the ratio of the weight of a given volume of wood to the weight of an equal volume of water. It can be calculated based on oven-dry weight, and volume on oven-dry, air-dry or green condition. The most common methods utilized to measure small wood samples relative density are gravimetric analyses (Smith, 1954; Ifju, 1969; Elliott, 1970) and x-ray densitometric techniques (Parker et aj_. , 1973; Parker and Jozsa, 1973). The latter method is preferred because it easily provides a continuous readout of intra-ring and intei—ring components of cross-sectional wood cores, such as earlywood, latewood and total growth increment widths and relative densities. Relative density of a given growth increment increases as the proportion of latewood increases and the tracheids become smaller and thicker-walled (Zobel and Talbert, 1984). In Douglas-fir, the average growth increment relative density is controlled mainly by the proportion of latewood tracheids, due to the fact that the relative density of such tracheids is between two and three times higher than that of earlywood tracheids (Ifju and Kennedy, 1962; Ifju eta].., 1965). 20 In some species, the variations in relative density are primarily correlated to the number of growth increments from the pith (age) rather than to the growth rate. This means that the growth rates and relative densities of trees of comparable environment, species, age and height are virtually independent traits (Turnbul1, 1937; Rendle and Phillips, 1957). In the case of Douglas-fir and many pine species, several studies have substantiated this trend (Goggans, 1961; Smith et aj_., 1966; McKimmy, 1966; Kennedy and Warren, 1969; De Guth, 1980; Pearson and GiImore, 1980; Barrett and Kellogg, 1984; Pearson and Ross, 1984). In spruce and fir, however, fast growth rate is usually associated with low relative density (Hale and Fenson, 1931; Hale and Prince, 1940; Aldridge and Hudson, 1959; Chang and Kennedy, 1967). Within tree relative density variation can be assessed in a horizontal, vertical or diagonal classification scheme (Duff and Nolan, 1953; Forward and Nolan, 1964). The horizontal scheme is the most studied and represents the horizontally or diametrically arranged sequence of growth increments from pith to bark. The vertical scheme represents the vertically arranged sequence of growth increments at the same cambial age. Finally, the diagonal scheme represents the diagonally arranged sequence of growth increments for the same calendar year. 21 According to the horizontal relative density variation, the juvenile wood relative density can be lower than, higher than or approximately the same as that of mature wood, depending upon species. The most common trend found is a low juvenile wood relative density which gradually increases from the pith to mature wood, where it stabilizes. This trend was described by Loo et al_. (1985) for loblolly pine, by Foelkel et al_. (1976) for slash pine, by Bower et aJL- (1976) for Caribbean pine, by Harris (1969a) for radiata pine, by Paul (1950), Wellwood (1952), Littleford (1961), Harris (1969a), Cown (1976), Gerhards (1979), and by Barrett and Kellogg (1984) for Douglas-fir. The juvenile wood relative density in Douglas-fir was also found highest near the pith, decreasing rapidly in the first growth increments from the pith, then increasing outward to the mature wood (Chalk, 1953; Harris and Orman, 1958; Kennedy and Warren, 1969; Cown, 1976; Megraw and Nearn, 1972; Jozsa and Kellogg, 1986). The juvenile wood relative density was found to be higher than that of mature wood by Boutelje (1968) for Norway spruce (Picea abi es (L.) Karst.), by Taylor et al. (1982), and Wang and Micko (1984) for white spruce, by Jozsa and Kellogg (1986) for interior white spruce or interior Engelmann spruce (Pi cea enge1mann i i Parry), by Wood and Bryan (1960) for Sitka spruce (Pi cea s i tchens i s 22 (Bong.) Carr.), by Polge (1964), Well wood and Jurazs (1968), and Jozsa and Kellogg (1986) for western redcedar, and by Well wood and Smith (1962), Krahmer (1966), and Jozsa and Kellogg (1986) for western hemlock (Tsuga heterophylla (Raf.) Sarg.). Little or no horizontal relative density variation between juvenile and mature wood was found by Harris (1969a) for balsam fir (Abies balsamea (L.) Mill.) and Norway spruce, and by Jozsa and Kellogg (1986) for lodgepole pine (Pinus contorta Dougl.) and interior Douglas-fir (Pseudotsuga menz i es i i var. glauca (Beissn.) Franco). Considering the vertical relative density variation scheme, the juvenile (crown-formed) wood relative density was reported as significantly lower than that for mature (stem-formed) wood by Cooper (1960) and Brunden (1964) for red pine (Pinus resinosa Ait.). Well wood (1952, 1960) studying the density variation of 130 Douglas-fir and 39 western hemlock, sampled at stump height, one-third total height and the top of the merchantable stem, found that relative density decreased significantly with increased height of the trees. In contrast, Polge (1964) and Wei 1 wood and Jurazs (1968) found that the crown-formed wood relative density was higher than that of the stem-formed wood in western redcedar. 23 A slight vertical relative density variation, probably associated with the presence of compression wood in the growth increments close to the pith, has been found within the juvenile wood core of some species. For instance, a small density increase in the terminal portion of the live crown was reported by Harris and Orman (1958) and Kellogg and Kennedy (1986) for Douglas-fir, by Zobel et aj_. (1959) for loblolly pine and slash pine, and by Krahmer (1966) for western hemlock. Finally, the diagonal relative density variation scheme is characterized by an initial decrease from the apex to approximately the limit of the live crown, followed by an increase toward the stem base. This trend can be seen from the data of Harris and Orman (1958) on Douglas-fir and Richardson (1961) on Corsican pine (Pinus  nigra var. mar itima (Alt.) Melville). The most commonly found relative density variation patterns in Douglas-fir are summarized below. 1. Relative density variation patterns are primarily related to the proportion of the latewood tracheids and to the number of growth increments from the pith. 2. Relative density in juvenile wood is high near the pith and in the terminal portion of the live crown, decreases rapidly in the 24 first growth increments from the pith and increases outward toward the mature wood, where it stabilizes. 3. In interior Douglas-fir there is a slight or no relative density variation from juvenile to mature wood. 2.2.2.2 Other Properties The physical properties of strength, elasticity and moisture content, in combination with the presence of knots and growth related defects, such as compression wood and spiral grain, determine the suitability of juvenile and mature wood for several end uses. The variations of the strength and elastic properties of juvenile and mature wood are mainly correlated to the respective variations in relative density. In general, the strength and elasticity of dimension lumber manufactured completely from, or containing a high percentage of, juvenile wood, has been found to be less than the strength and elasticity of mature wood. This trend was reported by Boone and Chudnoff (1972) and Bower et al. (1976) for Caribbean pine, by Pearson and Gilmore (1971, 1980) and Pearson and Ross (1984) for loblolly pine, and by Wangaard and Zumwalt (1949), Littleford (1961), 25 Gerhards (1979), Barrett and Kellogg (1984) and Senft et al. (1986) for Douglas-fir. Juvenile wood moisture content has been reported to be higher than that of mature wood by Zobel et a]_. (1968) and Zobel et aj_. (1972) for loblolly and slash pine, and by Britt (1970) for loblolly pine. Compression wood in the juvenile wood was reported by Boone and Chudnoff (1972) for Caribbean pine, by Crist et a 1. (1977) for jack pine (Pinus banksiana Lamb.) and eastern larch, and by Zobel and Kellison (1972) and Zobel et al. (1972) for loblolly and slash pines. In Douglas-fir, the possible presence of compression wood in juvenile wood has been suggested as the principle cause of the high density values observed in the growth increments close to the pith (Chalk, 1953; Harris and Orman, 1958; Kennedy and Warren, 1969; Kellogg and Kennedy, 1986). High spiral grain angles in Juvenile wood were reported by Harris (1969b) for radiata pine and by Zobel et al. (1972) for slash and loblolly pines. In Douglas-fir, Northcott (1957), Elliott (1958) and Woodfin (1969) observed that the most frequent spiral pattern was in the left direction near the pith, with the angle increasing in the first formed growth increments in the juvenile wood, and then changing gradually from left to right spiralfty as the tree age increased. 26 2.2.3 Juvenile - Mature Wood Transition Determination Determination of the transition zone from juvenile to mature wood is critical for setting relative proportions, and important in definition of wood quality and timber value. This determination can be difficult, however, because as juvenile wood matures, the changes in its properties are gradual and often erratic. The characteristics of change differ according to species and wood properties selected for analysis. For example, the mechanical properties and relative density of wood generally indicate maturity earlier than tracheid length and microfibril angle (Bendtsen, 1978; Megraw, 1985; Bendtsen and Senft, 1986). Some species of spruce (Pi cea spp.), fir (Abi es spp.) and cypress (Cuppressus spp.) are characterized by an indistinct juvenile - mature wood transition zone. Therefore, the differences in properties between juvenile and mature wood are difficult to observe (Zobel, 1984). In contrast, Douglas-fir and most hard pines, such as loblolly pine, slash pine and Caribbean pine, show a more distinct juvenile - mature wood transition zone. 27 Several subjective and objective methods have been utilized to define the transition zone between juvenile and mature wood. 2.2.3.1 Subjective Methods The first, and perhaps the most general, method of determining the juvenile - mature wood transition zone from the subjective viewpoint is visual examination of stem cross sections or increments cores. For instance, Zobel et al. (1959) defined the transition zone for slash pine as the first five to eight growth increments from the pith, and for loblolly pine as the first seven to 11 growth increments from the pith, based on the dull and lifeless appearance of juvenile wood stem cross section samples when dr i ed. Yang et a±. (1986) defined the transition zone in eastern larch (Larix 1aricina (Du Roi) K. Koch) as the first four to 45 growth increments from the pith, depending on the sampling height. The transition zone was visually determined in two cm wide diametric strip wood samples based on the light colour and width of growth increments in the juvenile zone. 28 Other authors have defined the transition zone by comparing the wood properties of young trees with the wood properties of old trees. Zobel and Kellison (1972) used this method when defining the transition zone as the first ten growth increments from the pith, by comparing the wood properties of 11-year-old juvenile to 30-year-old mature loblolly pine trees. Boone and Chudnoff (1972) and Bower et al. (1976) utilized the same approach when comparing plantation grown and forest grown Caribbean pine trees. The transition zone can also be defined by comparing the wood properties of juvenile (crown-formed) wood to mature (stem-formed) wood. The age at the base of the live crown can then be identified as the transition age between juvenile and mature wood. Brunden (1964), studying red pine, found that the length and relative density of stem-formed tracheids were significantly higher than those from crown-formed wood. He did not, however, consider the age at the base of the live crown. Cooper (1960) found that red pine stem-formed wood relative density was significantly higher than that of the crown-formed wood. He identified the age at the base of the live crown as approximately 20 years. Finally, the juvenile - mature wood transition zone has been determined on the basis of changes in certain anatomical or physical wood properties between the closest 29 and farthest growth increments from the pith. The transition zone is defined as the location where the values of these properties begin to remain constant. In southern and tropical pines, the transition zone ranges from six to 15 growth increments from the pith (Zobel and McElwee, 1958; Pearson and Gil more, 1980; Bendtsen, 1978; Zobel, 1984; Senft et al., 1985). In Japanese larch (Larix  1epto1ep i s (Sieb. and Zucc.) Endl.) and white spruce, the transition zone has been reported as the first ten growth increments from the pith by Isebrands and Hunt (1975) and by Taylor et aj_. (1982). In Douglas-fir, the transition zone has been reported as the first 15 to 20 growth increments from the pith (Wellwood and Smith, 1962; McKimmy, 1966; Erickson and Arima, 1974; Erickson and Harrison, 1974; McKimmy and Campbell, 1982; Barrett and Kellogg, 1984; Senft et al•, 1985; Jackson and Megraw, 1986; Senft et aj_. , 1986; Jozsa and Kellogg, 1986). 2.2.3.2 Objective Methods Recently, some researchers have utilized more objective methods to define the zone of transition between juvenile and mature wood. For instance, Shiokura (1984) used a logarithmic regression equation to relate tracheid 30 length to the number of growth increments from the pith (age). The transition zone was defined when a one percent tracheid length difference was recorded between two consecutive growth increments. The transition ages at different heights were 11 to 19 on Japanese larch, 14 to 18 on Sakhalin fir (Abi es sacha1i nens i s Mast.) and on Hondo spruce (Pi cea Jezoens i s (Sieb. and Zucc.) Carr.) and 20 to 22 on Japanese red cedar (Cryptomeria Japonica (L.F.) Don). Yang et al_. (1986) utilized two simple linear regression models to relate tracheid length to number of growth increments from the pith. The first model was fitted in the juvenile wood zone, where the length of tracheids increased, and the second was fitted in the mature wood zone, where the length of the tracheids remained constant. The transition zone, which was determined as the age at which the juvenile and mature regression models intersected, was found to be from ten to 44, depending on tree height. Loo et aj_. (1985), studying the genetic variation in the time of transition from juvenile to mature wood in loblolly pine, utilized a combination of three methods to define the transition age, and considered the properties of specific gravity and tracheid length. The first method involved fitting two simple linear regression models to the data, which were plotted as functions of age. The age of 31 the transition was estimated as the age at which the second model (mature wood) showed the best fit, determined by the smallest residual sum of squares. The second method was used when the best fitted mature wood model, using the first method, showed a negative slope. In this case, the slope of the mature wood model was held constant at zero, assuming that the mature wood values fluctuate around the constant mean. The third method was a visual examination, used when the data points did not conform to the former two patterns of variation. The mean ages of transition were found to be 11.45 and 10.30 years for specific gravity and tracheid length, respectively. Bendtsen and Senft (1986) applied three methods to both individual tree and average values of strength, elasticity, specific gravity, cell length and fibril angle to determine the transition ages for loblolly pine and cottonwood (Popu1 us de1 toides Bartr.). The methods used were segmented regression analysis, discriminate analysis, and analysis of slope. None of these methods produced a consistent demarcation between juvenile and mature wood because of the large variability among values from tree to tree and year to year. Therefore, visual interpretations of the data and data plots were used to determine the transition ages, which were, depending on the property analyzed, 12 to 18 for loblolly pine and 17 to 18 for cottonwood. 32 In conclusion, it can be said that the age of the juvenile - mature wood transition ranges from five to 18 years in southern and tropical pines, from 15 to 20 years in Douglas-fir, and from four to 44 years in the rest of the species described, depending on the sampling height and wood property analyzed. In some species of spruce, fir and cypress, the juvenile - mature wood transition zone is not clearly defined. 33 3.0 MATERIALS AND METHODS In order to test the project hypothesis, 13 unpruned, two pruned second-growth, and two unpruned plantation-grown coastal Douglas-fir trees were sampled from nine stands as fo11ows. Six second-growth stands were selected on the west and east coasts of Vancouver Island, British Columbia as a part of the Douglas-fir Task Force Project (Kellogg, 1986). The stands were approximately 50 years old and growing on sites of medium to good productivity. Ten dominant and codominant Douglas-fir trees of uniform growth rate (growth increments of approximately four to five mm per year) were selected from each stand by the Task Force for basic wood property studies. The two felled trees per stand with the most intact crowns were selected for the present study of the transition from juvenile wood to mature wood. Five additional trees were selected to extend sampling to pruned stands, and to sites of different productivity. One tree was measured in each of two 37-year-old plantations growing on soil of high (marine clay) and low (gravel outwash) productivity located west of Campbell River. One unpruned 21 yeai—old tree, and two pruned trees of 51 and 45 years of age were sampled from the University of British Columbia 34 Research Forest at Haney. The pruned trees were approximately 15 cm D.B.H. and approximately 14 m tall when the lower one-third of the live crown was removed to a height of four to five metres in 1954. The location of the stands is shown in Figure 1, while stand characteristics are given in Table 1. Sample tree data are shown in Table 2. 3.1 External Tree Characteristics The external tree characteristics were categorized as stem and crown respectively. The following stem measurements were made and recorded: 1. Age at stump; 2. Total tree height; 3. Internodal distances over the length of the stem; 4. Height to base of live crown (distance from the ground to the lowest whorl which has three or more live branches); and 5. Inside and outside bark diameters of five cross-sectional discs, eight to 16 cm thick, cut from breast height, and positions 20, 40, 60 and 80 percent of total tree height. 35 These measurements were taken to estimate the curve representing cumulative height growth over number of growth increments from the pith for each tree. The cross-sectional discs were taken to Forintek Canada Corporation's x-ray densitometry laboratory in Vancouver for further analysis. To study the actual crown characteristics, crown cover, live branches and height to base of the live crown were measured as follows. The crown cover, or ground area covered by the vertical projection of each crown, was measured using a metric tape and a Suunto clinometer with a 90 degree scale (Husch et aj_. , 1982). An average crown radius was estimated from six to seven crown radii measurements taken every 45 degrees around the perimeter of the crown and towards competing trees (Table 3). Length and height position of two live branches of similar diameter from opposite sides of every second whorl were measured from each tree. The height to base of live crown was determined by measuring the distance from the ground to the lowest whorl which had three or more live branches. In addition, an average crown height was calculated as a function of the average crown radius, which represented the maximum length of the branches located at the widest part of the crown. Average crown height was estimated in order to relate the results of this project to 36 the Ministry of Forests and Lands Tree and Stand Simulator Model (TASS) (Mitchell, 1975, 1980; Mitchell and Cameron, 1985). The procedure for estimating average crown height is summarized in Appendix 1 and the results are contained i n Tab 1e 3. To study the development of the crown over time, two representative dead branches or branch stubs were collected from every second or third branch whorl. Each branch was examined to determine the number of growth increments at the base, which represented the age at which the branch stopped producing growth increments. Discs of one cm thickness were cut from each branch base, labelled and sanded. The growth increments in each branch disc were counted and recorded using a low power stereoscopic microscope. The number of growth increments in each branch disc and the corresponding whorl height position were utilized to determine the approximate position of the base of the live crown at younger ages. An example of this determination is shown in Appendix 2. 3.2 Internal Tree Characteristics To study the internal tree characteristics, x-ray densitometric techniques were utilized to determine growth increment relative density data. Linear regression 37 techniques were utilized to estimate the juvenile - mature wood transition age from the data. 3.2.1 X-Ray Densitometric Analyses For the x-ray densitometric evaluation, a total of 85 cross-sectional discs (17 trees, five sample heights) of eight to 16 cm thick, were sampled. On each disc, two average radii sections at least 90 degrees apart were measured and labelled, avoiding knots, resin pockets and compression wood. The average radii were first cut into strips of one cm thick and 10 cm wide. They were then re-cut into smaller sample strips, five mm wide in the tangential direction and six mm thick along the grain, resembling increment cores. The wood sample strips were utilized for x-ray densitometr1c analysis following the procedures explained in detail by Parker and Jozsa (1973) and by Parker et aj_. (1973, 1980). A summary of these procedures is presented in Appendix 3. A total of 170 wood core radii were scanned using the above procedure. The relative density values were expressed on an oven-dry volume and weight basis. Annual pith to bark relative density profiles (average relative density of two core radii growth increments per sample section) were plotted to determine the annual growth 38 increment relative density variation patterns as a function of number of growth increments from the pith and height position on the stem of each sample tree, as shown in Figure 2, for Sample Tree 1A5. 3.2.2 Juvenile - Mature Wood Transition Determination After plotting the relative density profiles of each sample section, a visual examination was done to detect the lowest relative density value and a possible juvenile -mature wood zone based on the shape of the profile. Two categories of profiles were identified. The first category included the relative density profiles which showed fairly constant relative density values from the pith to bark, without a clear definition of transition zone. In this case, a simple linear regression model was fitted to each profile, starting from the lowest relative density value outwards, and it was assumed that the profile represented juvenile wood entirely. The second category included the profiles that showed an initial decrease in relative density in the first growth increments from the pith, followed by a gradual linear increase for a certain number of increments, then a final levelling off outwards. This relative density variation pattern indicated the possible evidence of a transition 39 zone in the profile. To determine such transition zone, a segmented regression analysis of each relative density profile was done as described below (Hudson, 1966; Draper and Smith, 1981). 1. A segmented regression model, which combines two linear segments, was fitted to each of the profiles starting at the lowest relative density value. The model utilized was: y = bQ + bjb2 + b3(x - b2> where: y = predicted value (relative density) bg = intercept of first line segment bj = slope of first line segment t>2 - transition age estimate bg = slope of second line segment x = independent variable (number of growth increments from the pith) 2. The best fitted model was determined by minimizing the total residual sum of squares of every possible division of the points between the first and the second segment. This method is called the Least Squares Line of Best Fit and is given by: 40 n n I e2f = I ( yf - y, ) 2 i = 1 i = l where: n 2 £ e j = residual sum of squares i = 1 y^ = observed values (relative density) A y. = predicted values (relative density) The division and set of estimates that gave rise to the smallest residual sum of squares was selected. This was done by means of a Fortran program which included non-linear optimization routines (Appendix 4). 3. The constraints of this procedure were that the first segment must start at the lowest relative density data point and that the intersection point between the two segments must be between two consecutive growth increments that split the data set. 4. The point of intersection between the two linear segments was identified to determine the number of growth increments (age) at which the transition from juvenile to mature wood occurred. 41 To verify the need for a segmented regression analysis, a statistical "F" test was done, calculating the ratio between the mean sum of squares residual value of a simple linear regression model and that of a segmented regression model, both fitted to the same profile (Figures 3a and 3b). When the "F" ratios calculated were greater than the "F" values tabulated by Pearson and Hartley (1954) the segmented regression model was chosen to determine the juvenile - mature wood transition zone. When the "F" ratios calculated were lower than the "F" values tabulated, the simple regression model was chosen and the juvenile - mature wood transition zone could not be determined. In this case, it was assumed that the relative density profile represented juvenile wood ent i re 1y. The determinations of the transition zone by the segmented regression analysis were checked against the corresponding profile to ensure that the regression model used was the most reasonable choice with respect to the data. 42 After determination of the juvenile - mature wood transition age for each relative density profile, relationships between number of growth increments from the pith and tree height position of both transition zone and base of the live crown were established. These within tree and between tree relationships were utilized to test the project hypothesis, and can be used to further extend the TASS model (Mitchell, 1975, 1980; Mitchell and Cameron, 1985) for predicting the volume of both juvenile and mature wood. In addition, average juvenile and mature annual growth Increment relative densities were determined to observe the within tree and between tree relative density var iat ions. 4.0 RESULTS AND DISCUSSION Table 2 gives a summary of sample tree characteristics. The plantation-grown and second-growth trees were dominant and codominant with ages at breast height ranging from 21 to 63 years, diameters at breast height from 19.8 to 68.2 cm and total heights from 16.8 to 44.7 m. These plantation-grown and second-growth trees were analyzed separately from the pruned trees. It must be noted that sample trees were selected from the nine sites without attempting to achieve a representation of the total popu1 at i on. 4.1 X-Ray Densitometric Analyses Summaries of the relative density values of 15 trees for sections sampled at breast height, 20 percent, 40 percent, 60 percent and 80 percent of total tree height are presented in Tables 4 to 8 respectively. For each sample section, relative density is expressed as the mean of all annual growth increment means. Pith to bark relative density profiles are illustrated for Sample Tree 1A5 in Figure 2. 44 The general relative density variation pattern of all sections sampled at breast height showed an initial decrease from the pith to a point within the second through tenth growth increments followed by a linear increase which gradually stabilized outwards over subsequent growth increments from the pith. Overall mean relative density for these sections was 0.523, with a maximum of 0.625 and a minimum of 0.413. The number of growth increments varied from 21 to 63 (Table 4). At 20 percent of total height the relative density variation pattern was similar to that of all sections at breast height. Overal1 mean relative density was 0.485, with a maximum of 0.589 and a minimum of 0.389. The number of growth Increments varied from 17 to 53 and sample height from 3.3 to 8.5m (Tab1e 5). Sections sampled at 40 percent of total height showed an initial decline in relative density from the pith to a point within the third through fifteenth growth increments, followed by a gradual increase outwards. In a few cases, a gradual decrease outwards was observed also. Overall mean relative density for these sections was 0.467, with a maximum of 0.549 and a minimum of 0.400. The number of growth increments ranged from 13 to 45 and sample height from 5.1 to 17.2 m (Tab 1e 6). 45 At 60 percent of total height there was an initial decline in relative density from the pith to a point within the fourth through fourteenth growth increments, followed by a moderate increase outwards. Overall mean relative density for these sections was 0.462 with a maximum of 0.551 and a minimum of 0.410. The number of growth increments varied from eight to 31 and sample height from 9.9 to 25.8 m (Table 7). The sections sampled at 80 percent of total height showed an initial decrease in relative density from the pith to a point within the fourth through twelfth growth increments, followed by an irregular and moderate increase outwards. Overall mean relative density for these sections was 0.478 with a maximum of 0.556 and a minimum of 0.437. The number of growth increments varied from four to 19 and sample height varied from 13.4 to 36.1 m (Table 8). From the above results, it would appear that the horizontal relative density variation pattern at the five sample heights is characterized by an initial decrease from the pith followed by a gradual increase and stabilization outwards, particularly in the lower sections (i.e. breast height, 20 and 40 percent of total height). Similar horizontal relative density variation patterns for Douglas-fir trees were reported by Paul (1950), Wei 1 wood (1952), Chalk (1953), Littleford (1961), Harris (1969a), 46 Kennedy and Warren (1969), Cown (1976), Gerhards (1979), Barrett and Kellogg (1984), and by Jozsa and Kellogg (1986). The vertical overall mean relative density for sections at breast height to 60 percent of total height (Tables 4 to 8) showed a decrease from 0.523 to 0.462, followed by an increase to 0.478 for sections at 80 percent of total height. In percentages, the decrease in relative density from breast height (100 percent) represented approximately seven, 10, 12 and eight percent respectively. This indicates that, in general, relative density decreases with height of sampling. Qualitatively similar density vertical variation patterns were reported by We11 wood (1952, 1960), and by Megraw (1986). 4.2 Juvenile - Mature Wood Transition Determination A total of 75 relative density profiles (15 trees, five sample height positions per tree) were investigated in order to detect transition zones between juvenile and mature wood. The profiles were divided into two categories based on the configuration of the data points pertaining to relative density and number of growth increments from the Pith. 47 The first category involved a total of 38 profiles characterized by having no clear definition of a juvenile -mature wood transition zone. In general, these profiles showed an initial decrease in relative density from the pith to a point within the fourth through thirteenth growth increments, followed by a gradual increase, decrease or levelling off outwards. To illustrate such trends, simple linear regression models were fitted to each profile, starting from the lowest relative density value outwards, assuming that they represented juvenile wood entirely (Appendices 5a to 19a). Table 9 shows a summary of the distribution and characteristics of these profiles among sample heights. The average number of growth increments was 18.42, ranging from 4 to 33, and the average section height was 20.99 m, ranging from 1.30 to 36.10 m. It can be seen that about 80 percent of the cases for which no evidence of juvenile -mature wood transition zone was found were at sections 60 and 80 percent of total height. In 15 percent of the cases no transition was found at 40 percent of total height. Only in five percent of the cases was no transition found at sections sampled at breast height and 20 percent of total height, corresponding to the younger sampled tree, i.e. RF1 (Appendix 19). 48 From the above, it is concluded that mature wood and therefore the juvenile - mature wood transition zone, was not present in the upper 40 percent of the stem nor in lower sections containing few growth increments from the pith (i.e. 13 to 33). The second category comprises relative density profiles which gave evidence of a transition zone. These profiles, in general, showed an initial relative density decrease in the first growth increments from the pith, followed by a gradual increase over several additional growth increments, then a final levelling off outwards. A total of 37 profiles from 14 trees were investigated in this section. Segmented linear regression models, starting at the lowest relative density value, were fitted to these profiles in order to determine the juvenile -mature wood transition zone (Appendices 5a to 18a). The point of intersection between the two linear segments corresponding to the segmented model was identified as the number of growth increments from the pith (age) at which the transition from juvenile to mature wood occurred. Table 10 gives a summary of the transition ages of each tree sample section, determined using segmented linear regression models. To verify the need of such models, the "F" ratio of the mean sum of squares residual value of a 49 simple linear regression model to that of a segmented model, both fitted to each relative density profile, was calculated for each sample section (Table 10). In al1 cases, the calculated "F" ratios were greater than one, indicating that the segmented regression model showed a smaller sum of squares residual value, thus a better data fit. The probability level at which the calculated "F" ratios were statistically significant varied from 0.001 to 0.470. Despite such a large range of level of significance, it was assumed that the segmented model was the best representation of the high variability among the values in some data sets, particularly considering that 56 percent of the "F" ratios were within the 0.30 level of probabi1ity. Figure 4a illustrates the distribution of sample section heights over the number of growth increments from the pith (age) at which the transition occurred, when segmented regression models were fitted to the relative density profiles. As can be seen, no systematic variation trend was found, indicating that transition age and sample height are independent. Table 11 gives a summary of average transition ages by sample section heights. These averages were 22.36, 22.86, 20.89 and 22.18 years for sections sampled at breast height, 20 percent, 40 percent of total height and for all sections, respectively. The 50 similarity of the average transition ages also indicates that transition age and height of sampling are not related. In addition, the average transition age of 22.18 years for all sections was very close to the maximum reported transition age for Douglas-fir, which is 20 years (McKimmy and Campbell, 1982; Barrett and Kellogg, 1984; Senft et. aj.. , 1985; Jozsa and Swan, 1986). 4.2.1 Juvenile - Mature Wood Relative Density Values Annual juvenile and mature wood relative density values were determined to observe within tree and between tree relative density variations. Summaries of relative density values for each tree are presented in Appendices 20a to 20o. Table 12 summarizes the relative density profiles which illustrated no clear juvenile - mature wood transition zone. These profiles, considered as representative of juvenile wood, had an average number of growth increments from the pith of 18.42 and an overall mean relative density of 0.467 ranging from 0.416 to 0.551. Table 13 summarizes the relative density profiles which showed a defined juvenile - mature wood transition zone. For these profiles, the average numbers of growth increments contained within the juvenile and mature wood were 22.18 and 19.95, respectively. All sample sections 51 mean relative densities were 0.472, 0.533 and 0.501 for juvenile, mature and total respectively. Table 14 summarizes relative density values for juvenile and mature wood. This is a combination of the results shown in Tables 12 and 13. The overall average numbers of growth increments representing juvenile and mature wood were 20.28 and 19.95, respectively. The juvenile wood transition value is almost identical to the reported maximum transition value of 20 growth increments. The overall mean juvenile wood relative density was 0.469, ranging from 0.410 to 0.561. The lowest juvenile wood relative density values were found at sections sampled at 20 and 40 percent of tree height. The overall mean mature wood relative density was 0.533 (14 percent greater than that for juvenile wood), ranging from 0.474 to 0.594. 4.3 Base of the Live Crown A total of 120 dead branches were sampled from 12 trees in order to approximate individual height to live crown base curves before harvest, as illustrated in Appendix 2. Table 15 shows a summary of the average number, height position, and number of growth increments found at the base of dead branches in each tree. Figure 4b 52 illustrates the overall distribution of the branches among trees as a function of height position in stem and number of growth increments at the branch base. As can be seen, the general trend was that the number of growth increments found in dead branches increased with tree height. It is Important to note that as the branches get older, their vigor decreases, leading to the formation of incomplete or absent growth increments at the branch base (Andrew and Gill, 1939; Reukema, 1959, 1961). Although few live branches located at the crown base were examined, the tendency was to find some discrepancies between the number of growth increments at the branch base and that of the tree stem at the point of intersection. Growth increment, counts showed five to eight fewer growth increments in the lower crown branches than in the stems at points of i ntersect i on. Reukema (1959) stated two possible reasons for this apparent failure of branches in forming growth increments. The first was that the cell formation ceased during the latter years of the branch's life. The second was that a considerable decrease in cambial activity at this time produced very narrow growth increments, i.e. one to two cells wide, without ear 1ywood and latewood differentiation. 53 From the above, it can be concluded that despite the fact that the lower crown branches appeared healthy, with plentiful foliage, they are probably no longer part of the functional crown. This may introduce some possibility of error when the base of the live crown is reconstructed based upon the number of growth increments found at the base of dead branches. In addition, this represents an argument in favor of pruning lower crown live branches. Table 3 summarizes the crown characteristics measured for the sampled trees at time of harvest. As illustrated, the overall crown base height, estimated as the height from the ground to the lowest whorl at which three or more live branches were located, was 19.7 m, ranging from 9.0 to 24.8 m. The overal1 average crown height, estimated as a function of the average crown radius (Appendix 1) was 24.2 m, ranging from 12.0 to 35.0 m. In some cases, this position was also Identified as the crown contact or point of intersection with neighbouring trees. This was done by looking at symptoms of whipping or breaking damage on the terminal shoots of the branches. Figures 5a and 5b illustrate the observed data distribution and height prediction linear regression models for crown base height and average crown height as functions 54 of total tree height. Figure 6 shows the former linear regression models with the addition of the observed total tree heights for each tree. As can be seen, both crown base height and average crown height increase with increasing total tree height at a very constant rate of change. This indicates a fairly consistent within and between crown measurement variation as a function of total height. 4.4 Relationships Between Juvenile - Mature Wood Transition and Base of the Live Crown A series of graphical representations were prepared in order to clarify the project hypothesis. Figure 7 shows two hypothetical curves representing total tree height and height to crown base over the number of growth increments from the pith at breast height. If the project hypothesis is correct, the curve representing height to crown base will also represent the points of juvenile - mature wood transition occurring at different heights and ages in the tree. Alternatively, if the hypothesis is not correct, the height to crown base will not be coincident with the points at which the juvenile - mature wood transition occurs. Figure 8 illustrates the height to crown base before harvest, the observed base of the live crown and calculated 55 average crown height, i.e. distance from the ground to the widest part of the crown, at harvest. The next step is to relate the former crown height positions to the corresponding juvenile - mature wood transition points. These relationships can, then, be established before or after harvest in order to test the project hypothesis as follows. 4.4.1 Hypothesis Testing Before Harvest Appendices 5b to 19b illustrate individual and overall (Figures 9a, 10a and 10b) relationships for total height, height to crown base and height to juvenile - mature wood transition points as functions of number of growth increments from the pith at breast height. In addition, the height to the lowest relative density values in each sample section was also plotted to establish relationships with the former curves (Figure 9b and Appendices 5b to 19b). The lowest relative density values presented in Figure 4c and Table 16, were the starting points for the segmented and simple linear models fitted to each profile. From Appendices 5b to 19b, the following individual tree characteristics are summarized below. 1. The curves representing height over number of growth increments from the pith at breast 56 height were very similar and can be classified within Site Classes I and II (i.e. good and medium good) for coastal Douglas-fir, according to Bruce (1981). The exception was tree CR1 (Appendix 17b) which was sampled from low productivity Site Class IV (i.e. med i um poor). The lowest relative density curves were, in general, lower and parallel to the total height curves. The average difference between the number of growth increments between these curves was 7.1, with a range of two to 15 and a standard deviation of 2.9 (Table 16). This suggests not only a very consistent pattern of decreasing relative density in the first growth increments from the pith, but also a tendency to find the lowest relative density value within a constant short distance from the tree apex at any height of sampling. This distance probably represents the upper one-third of the 1i ve crown. The height to crown base curves were parallel to the total height curves up to the region between 20 and 40 percent of the tree height, 57 and then they diverged towards the direction of the live crown base at harvest. This is to be expected, since the number of growth increments in branches increases with tree height up to the base of the live crown, and then decreases towards the apex. 4. The height to crown base curves were also parallel and sometimes coincident with the lowest relative density curves, particularly at lower sample heights. Therefore, it would appear that there is a biological connection between the number of growth increments found at the base of dead branches and that at which the annual increment relative density value is lowest. This would suggest that the productivity of a branch decreases with age, due to the declining photosynthetic efficiency and retention of needles. Growth increment relative density decreases proportionally until reaching a minimum value when the branch stops producing growth i ncrements. 5. The data points representing height position and number of growth increments from the pith at which the juvenile - mature wood 58 •transition occurs were lower and generally parallel to the former three curves (i.e. total height, lowest relative density, and crown base) for equal numbers of growth increments from the pith. The height difference between the height to crown base and the transition points slightly decreased with increasing sample height. This suggests that juvenile - mature wood transition occurs below the total height curve, the lowest relative density curve and the crown base curve, with a moderate tendency to approach the crown base curve at higher sample hei ghts. To illustrate general height variation trends, data from twelve trees were utilized to estimate the following overall height prediction models: y = 1.0909x°-9633 0.9968X y = tota1 he i ght x = growth increments from pith r = 0.9781 (correlation coefficient; 60 observations) y = -3.5855 + 0.8954x - 0.0037x2 y = lowest relative density height x = growth increments from pith r = 0.9324 (correlation coefficient; 60 observations) 59 y = -6.3482 + 1.0073x - 0.0088x y = crown base height x = growth increments from pith r = 0.9089 (correlation coefficient; 120 observations) y = -6.9988 + 0.4853x y = juvenile - mature wood transition height x = growth increments from pith r = 0.7174 (correlation coefficient; 37 observations) Trees CR1, CR2 and RF1 were not included in these models because they lacked either dead branch measurements or juvenile - mature wood transition determination. Figures 9a, 9b, 10a and 10b illustrate the observed data points and corresponding height prediction models for total height, lowest relative density height, crown base height and juvenile - mature wood transition height as a function of number of growth increments from the pith at breast height. Figure 11 combines the former four height prediction models with the addition of diagrammatic representations of trees at different states of development, to facilitate the interpretation of the results. It can be seen that the overall models follow the same variation patterns as those already described for individual observations on each tree. The lowest relative density curve was parallel and very close to the total height curve. This suggests that a minimum growth increment relative density value is likely 60 to be found within a short distance from the tree apex. The crown base curve was parallel to the total height curve up to about 40 percent of the total height where it started to diverge towards the position of the live crown base at harvest. This curve was also parallel and very closely related to the lowest relative density curve, particularly at lower sample heights. The simple linear regression model fitted to the points representing juvenile - mature wood transition was lower and parallel to the curves already described. However, due to the distribution of the data, this regression line does not accurately represent the mean values of the sections sampled at breast height and 40 percent of total height, as shown in Figure 10b. From the relative density point of view. Figure 11 can also be interpreted as a vertical relative density variation pattern. In this pattern, a maximum growth increment relative density value is found near the tree apex, a minimum value found within the upper one-third of the crown and a gradual increase in value found toward, and past, the crown base until the juvenile - mature wood transition zone, close to the base of the tree. The juvenile and mature growth increment mean relative density values reported in Table 14 support the interpretation of such a characteristic vertical relative density variation 61 pattern. Similar variation patterns were found by Chalk (1953) and by Harris and Orman (1958). The height difference between height to crown base and height to transition point, for equal numbers of growth increments from the pith, as a function of transition height, was plotted in Figure 12a. A definite downward trend with increasing transition height can be seen. Figure 12b illustrates the height difference between total tree height and height to transition point as a function of transition height. In this case, a slight downward trend with increasing transition height was also found, although the correlation coefficient was not significant at the 0.05 level of probability, i.e. r = -0.2531 (37 observations). Table 17 shows the average height differences between height of Juvenile - mature wood transition points, height to crown base and total height. The average differences between height of transition points and height to crown base were 10.25, 8.71 and 3.85 m for sections sampled at breast height, 20 and 40 percent of total height, respectively. The average differences between height to transition points and total height were 18.95, 18.51 and 14.59 m for sections sampled at breast height, 20 and 40 percent respectively. These average height differences confirm the variation trends shown in Figures 12a and 12b. When considering all sampled sections, the average juvenile 62 - mature wood transition height was found at 7.95 and 17.72 metres below the crown base and the tree apex (Table 17). From the above, since the transition did not occur at the base of the crown, it is concluded that the hypothesis, when analyzed before harvest, has not been proven. As discussed earlier, it is important to point out that discrepancies between the number of growth increments at the branch base and that of the tree stem at the point of intersection are likely to occur. Therefore, the height to crown base position at earlier ages, estimated by counting the number of growth increments at the base of the dead branches, can be overestimated. For instance, fewer growth increments in the lower crown branches than in the stem at points of intersection would define a higher position of the crown base, thus, a greater height difference between the transition point and the crown base. From the above, it is concluded that the differences between height of transition points and height of crown base could be reduced to a point at which the hypothesis could be proven, i.e. Juvenile - mature wood transition occurs at the base of the live crown. However, the extent of this reduction was not intensively investigated in the project. 63 4.4.2 Hypothesis Testing After Harvest The results of testing the project hypothesis after harvest are difficult to validate because a time gap will be required in order to confirm whether the transition between juvenile and mature wood occurs at the actual base of the live crown. However, a possible alternative is to estimate the occurrence of juvenile - mature wood transition heights as functions of earlier ages and heights at which the transition took place, i.e. before harvest. These height estimates, then, are based upon the calculation of an average of transition ages, called average transition height A, and an average of transition heights from the tree apex, called average transition height B, as shown in Figure 13. Figure 14 illustrates the number of growth increments corresponding to the juvenile - mature wood transition, i.e. determined using segmented regression models, from sections sampled at breast height, 20 and 40 percent, and overall section averages (average of transition ages) for each tree. Figure 15 illustrates the differences between total heights and earlier juvenile - mature wood transition heights from sections sampled at breast height, 20 and 40 percent, and overall section averages (average transition height B) for each tree. 64 For each tree, the average transition age was converted to average transition height A by interpolation from the curve representing height over number of growth increments from the pith at breast height (Appendices 5b to 18b). Both average transition heights A and B were calculated because no systematic data variation trend within and between trees was detected, as shown in Figures 14 and 15. Observed data distribution and height prediction models for average transition heights A and B, crown base height and average height as functions of total tree height are shown in Figures 16a, 16b, 5a and 5b respectively. Figure 17 shows a combination of the former four linear regression models with the addition of the corresponding total tree height of each tree. It can be seen that the height positions pertaining to average crown, average transition A, live crown base and average transition B increased with increasing total tree height. The slopes of these models indicated relatively constant rates of height change. The average crown height regression line was located higher than those corresponding to the average transition height A, live crown base height and average transition height B. The average transition height A regression line was higher than that corresponding to average transition height B and to live crown base 65 height. Only one tree at the lower height end showed a live crown base higher than the average transition height. A, reversing the variation pattern established at the higher height end. The same relationships can be seen in the summary of average height differences presented in Table 18. The average difference between total height and average crown height, average transition height A, crown base height and average transition height B were 10.26, 12.63, 14.76 and 17.91 m, respectively. Since earlier results showed that the overall average of juvenile - mature transition height was found 17.72 m below the tree apex (Table 17), it can be stated that average transition height B, located 17.91 below the tree apex, would better represent the possible occurrence of transition from the tree apex at harvest. The average height difference between transition height A and average crown height was 2.16 m and between transition height A and live crown base height it was -2.21 m (Table 18). The correlation coefficients obtained between transition height A, average crown height and live crown base height were 0.7936 and 0.8419, respectively (Table 19). The average height difference between average transition height B and average crown height was -3.12 m, and between average transition height B and live crown base height it was -7.50 m (Table 18). The correlation 66 coefficients between average transition height B» average crown height and live crown base height were 0.5888 and 0.7340, respectively (Table 19). From the above, it can be summarized that average transition height A occurred below the average crown height and above the live crown base height. Average transition height B occurred below both the average crown height and the live crown base height. Neither of the calculated juvenile - mature wood average transition heights A nor B occurred at the base of the live crown position after harvest. Average transition height A seemed to be closer to the average crown height and the live crown base height than the average transition height B. However, assuming that the transition would be likely to occur below the base of the live crown position, as demonstrated earlier, the average transition height A might overestimate the height of the transition. Average transition height B, then, would better represent the possible occurrence of the transition after harvest. As illustrated in Table 17 and Figure 12a, the difference between height to live crown base and height to transition points before harvest decrease with increasing tree height. The average height differences were 10.25, 8.71 and 3.85 m for sections sampled at breast height, 20 and 40 percent of total tree height respectively (Table 67 17). Assuming that these height differences will continue to decrease with tree height, the average transition height B found at 3.12 m (Table 18) below the observed base of the live crown position seemed a very reasonable estimate of the occurrence of the transition after harvest. In view of the above, it is concluded that the average juvenile - mature wood transition height B, calculated as a function of earlier transition height positions, best represents an actual estimate of transition occurrence. Since neither of the estimated average transition heights A nor B occurred at the base of the live crown position, it follows that the hypothesis when analyzed after harvest is not proven. The similar results found when testing the hypothesis before and after harvest can be, in part, supported by Larson (1969) who stated that: "The major changes in wood formation and quality occur in the lower stem beneath the living crown or beneath the most active branches of the crown." In this project, the transition from juvenile to mature wood occurred below the arbitrary definition of live crown base, i.e. height to the lowest whorl which has three or more live branches. However, Larson's interpretations of the juvenile - mature wood transition, the extension of the living crown and location of the most active branches 68 in the crown, may not be biologically coincident with the results of this project. This suggests that a wrong definition of what is in reality the base of the live crown, as it relates to wood formation, may also lead to a wrong answer when testing the project hypothesis. 4.5 Pruned Trees As was stated in the literature review, pruning of live and vigorous lower branches in the crown can produce a faster change from juvenile to mature wood, therefore reducing the proportion of juvenile wood core in a given tree. In this project, two pruned trees were analyzed in order to investigate the effects of pruning on annual increment relative wood density. The results were then analyzed to determine their relationship to the project hypothes i s. Figures 18 and 19 show the relative density profiles as a function of number of growth increments from the pith. As shown, additional sample sections at ten percent of total tree height were included because they were located within the pruning area. This area comprised the lower one-third of the live crown, representing a height of approximately four to five metres, i.e. below the sample sections at 20 percent of total height. In these figures, 69 the arrows in the relative density profiles on the sections sampled at breast height and 10 percent of total height indicate the year in which the pruning was done, i.e. 1954 or 30 years before the trees were harvested. The relative density profiles of sections sampled at breast height and 10 percent of total tree height in tree RF2 showed a gradual initial increase or decrease in the first growth increments from the pith, followed by a sudden increase soon after the pruning was done, and a final gradual decrease outwards. The relative density profiles for the same sample sections in tree RF3 showed a similar variation pattern, until after pruning was done, and then they distinctively showed a final levelling off outwards. This suggests that relative density can be increased by pruning of lower crown 1i ve branches. The different relative density variation patterns of these trees after pruning was probably due to the fact that RF2 was more dominant and open grown than RF3. Therefore, RF2 was able to overcome the effects of pruning in a relatively short time. The numbers of growth increments from the pith, before pruning, at breast height and 10 percent of total height were 21 and 17 for tree RF2, and 15 and 13 for tree RF3. The overall average number of growth increments of the 70 former sections was 16.5. This average number also represents an estimate of the juvenile - mature transition occurrence, since it delimits two different relative density variation patterns from the pith. For both trees, the mean relative density for juvenile wood was 0.441, ranging from 0.370 to 0.490. The mean relative density for mature wood was 0.494 (nine percent greater than that for juvenile wood), ranging from 0.640 to 0.410 (Appendices 20p and 20q). When the overall average number of growth increments at which the transition occurred after pruning, i.e. 16.5, was compared to that corresponding to trees without pruning, i.e. 22.18 (Table 11), the difference was about 26 percent. From the above, it is concluded that pruning of lower crown live branches can produce both an increase in annual increment mean relative density and a decrease in the proportion of juvenile wood in Douglas-fir by accelerating the occurrence of juvenile - mature wood transition. Since only two trees were analyzed, these results should be treated as exploratory. However, they are good indicators of possible benefits in addition to obtaining more clear wood, when pruning lower crown live branches. Furthermore, assuming that these branches contribute little or nothing to stem wood formation, while utilizing tree resources, it 71 might be worthwhile to consider the pruning of young-growth Doug 1as-f i r. When relating these results to the project hypothesis, it would appear that pruning can accelerate the juvenile -mature wood transition, which otherwise would need more time in which to happen. Then, if in fact the transition occurs below the base of the live crown in unpruned trees, the transition could be shifted upwards by pruning lower crown live branches. In view of the above, it is concluded that when the project hypothesis was tested on pruned trees, the juvenile - mature wood transition would occur at the modified base of the live crown, which represents the upper limit of pruning height. Therefore, the hypothesis can be proven as correct. It is important to note that more research on pruned trees is needed for further development of these results. 72 5.0 CONCLUSIONS 1) The horizontal annual increment mean relative density variation pattern at the five sample heights was characterized by an initial decrease to a point within the first growth increments from the pith, followed by a gradual increase and stabilization outwards, particularly in the lower sample sections (i.e. breast height, 20 and 40 percent of total tree height). 2) The vertical annual increment mean relative density variation pattern was characterized by a decrease with increasing tree height. 3) Based on the variation pattern of the data points, pertaining to annual increment relative density and number of growth increments from the pith, no evidence of juvenile - mature transition zone was found primarily at sections sampled at 60 and 80 percent of total tree height. The average number of growth increments from the- pith for these sections was 18.42, ranging from 4 to 33. 4) A greater evidence of juvenile - mature wood transition zone was found at sections sampled at breast height, 20 73 and 40 percent of total tree height. The average numbers of growth increments at which juvenile - mature wood transition occurred, when segmented regression models were fitted to relative density profiles, were 22.36, 22.86, 20.89 and 22.18 for sections sampled at breast height, 20 percent, 40 percent of total height and for all sections, respectively. Segmented regression analysis, therefore, produced a fairly consistent demarcation between juvenile and mature wood, among sample heights, despite the large variability among the values in some of the relative dens i ty prof i1es. 5) The overall annual increment mean mature wood relative density was 14 percent higher than that for Juvenile wood. The lowest annual increment mean juvenile wood relative density was found at sections sampled at 20 and 40 percent of total tree height. 6) Individual and overall height relationships, before harvest, for total height, height to crown base, height to the lowest relative density value and height to juvenile - mature wood transition point as functions of number of growth increments from the pith indicated that the: 74 Lowest relative density curve was parallel and very close to the total height curve, suggesting that a minimum growth Increment relative density value is likely to be found in the first growth increments from the pith and within a short distance from the tree apex; Height to crown base curve was parallel to the total height curve up to about 40 percent of the total height where it started to diverge towards the live crown base position at harvest; Height to crown base curve was also parallel and sometimes coincident with the lowest relative density curve, indicating a possible biological connection between the number of growth increments at the base of dead branches and that at which the annual increment relative density value is lowest; and Simple linear model fitted to the points indicating juvenile - mature wood transition was lower and parallel to the curves described previously. 75 Observed data distribution and height prediction models, after harvest, for total height, average crown height, live crown base height, average transition height A (estimated as a function of earlier transition ages) and average transition height B (estimated as a function of earlier transition heights) indicated that: a. The average crown height positions were higher than the positions corresponding to average transition A, live crown base and average transition B; b. The average transition A height positions were located between those positions corresponding to the average crown and base of the live crown; c. The average transition B height positions were located below those positions corresponding to the average crown and the base of the live crown; and d. Assuming that the transition would be likely to occur below the live crown base position, the average transition height A might overestimate the height of the transition, then, the average transition height B would better represent the possible occurrence of the transition after harvest. 76 8) Pruning of lower crown live branches produced both an increase in the annual increment relative density and a decrease in the proportion of juvenile wood by accelerating the juvenile - mature wood transition. The average number of growth increments at which the transition occurred after pruning was 16.5. 9) When testing the project hypothesis on unpruned trees, before and after harvest, the transition in relative density from juvenile to mature wood did not occur at the base of the live crown, as defined. In both cases, the transition occurred below the base of the live crown. Therefore, the hypothesis has not been proven. 10) When the hypothesis was tested on pruned trees, the juvenile - mature wood transition did occur at the base of the live crown, which represented the upper limit of pruning height. The hypothesis in this case has, therefore, been proven. 11) Results of hypothesis testing seemed to depend upon the definition given to the base of the live crown. Further research needs to be done, not only to find a more precise definition of what should biologically be considered the base of the live crown, but also to investigate the implication of such definition on wood format ion. 77 LITERATURE CITED Aldridge, F. and R.H. Hudson. 1959. Growing quality of softwoods. Quart. J. Forestry 53:210-219. Allen, G.S. and J.N. Owens. 1972. The Life History of Douglas-fir. C.F.S. Cat. No. F042-4972. 139 pp. Anderson, E.A. 1951. Tracheid length variation in conifers as related to distance from pith. J. Forestry 49(l):38-42. Andrew, S.R. and L.S. Gill. 1939. Determining the time branches on living trees have been dead. J. Forestry 37:930-935. Barefoot, A.C, Hitchings, R.G. , El wood, E.L. and E. Wilson. 1970. The relationship between loblolly pine fiber morphology and kraft paper properties. Tech. Bull. No. 202. N.C Agric. Exp. St., N.C.S.U., Raleigh. 89 pp. Barrett, J.D. and R.M. Kellogg. 1984. Strength and stiffness of second-growth Douglas-fir dimension lumber. Forintek Canada Corp., Vancouver, B.C. Tech. Rep. 57 pp. . 1986. Lumber quality from second-growth managed forests. Proc. A Technical Workshop: What Does it Mean to Forest Management and Forest Products? F.P.R.S., S.A.F., Bellingham, Wa. pp. 57-71. Bendtsen, B.A. 1978. Properties of wood from improved and intensively managed trees. Forest Prod. J. 28(10):61-72. and J. Senft. 1986. Mechanical and anatomical properties in individual growth rings of plantation-grown eastern Cottonwood and loblolly pine. Wood and Fiber Sci. 18(l):23-38, Boone, R.S. and M. Chudnoff. 1972. Compression wood formation and other characteristics of plantation-grown Pi nus caribaea. U.S.D.A. For. Serv. Res. Pap. I.T.F.-13. Int. Trop. For., Puerto Rico. 15 pp. 78 Boutelje, J.B. 1968. Juvenile wood, with particular reference to northern spruce. Svensk Papptidn. 71(17):581-585. Bower, R.W., De Souza, A. and J.F. Senft. 1976. Physical and mechnical properties of fast-grown, plantation Caribbean pine (Pi nus. car ibaea) from Brazil, South America. Purdue Univ. Res. Bull. No 936. Lafayette, Ind. 6pp. Briggs, D.G. and W.R. Smith. 1986. Effects of silvicultural practices on wood properties of conifers - a review. Proc. of Douglas-fir Stand Management for the Future. Univ. Wa., Seattle, pp. 108-117. British Columbia Ministry of Forests. 1980. Forest and Range Resource Analysis. Technical Report. ISBN 0-7719-8324-7. Vol. II. pp. 577-602; 693-712. Britt, K.W. 1970. J_n Handbook of Pulp and Paper Techno1ogy. Zobe1, B.J. ed. Pu1pwood mensuration. Van Nostrand Reinhold Co. N.Y. pp.125-132. Brown, CL. 1970. Physiology of wood formation in conifers. Wood Sci. 3(l):8-22. Bruce, D.I. 1981. Consistent height-growth and growth rate estimates for remeasured plots. Forest Sci. 27(7):711-725. Brunden, M.N. 1964. Specific gravity and fiber length in crown-formed wood. Forest Prod. J. 14(1):13-17. Bryant, P.A.V. 1984. The impact of fast growth in plantations on wood quality and utilization. Proc. Symp. on Site and Productivity of Fast Growing Plantations. I.U.F.R.O., Pretoria, S. Africa. pp. 403-410. Chang, C.I. and R.W. Kennedy. 1967. Influence of specific gravity and growth rate on dry wood production in plantation grown white spruce. Forestry Chron. 43(2):165-173. Chalk, L. 1953. Variation of density in stems of Douglas-fir. Forestry 26:33-36. 79 Clark, J. 1961. Photosynthesis and respiration in white spruce and balsam fir. N.Y. State Coll. For., Syracuse Univ. Bull. 85. 72 pp. Cooper, G.A. 1960. Specific gravity of red pine as related to stem and crown-formed wood. Iowa State J. Sci . 34(4) -.693-708. Cown, D.J. 1973. Effect of severe thinning and pruning treatments on the intrinsic wood properties of young radiata pine. N.Z. J. For. Sci. 3(3):379-389. . 1975. Variations in tracheid dimensions in the stem of a 26-year-old radiata pine. Appita 28(4):237-245. . 1976. Densitometric studies on young Douglas-fir. Unpub. Ph. D. Thesis. Dept. of Forestry, Univ. B.C. 241 pp. Crist, J.B., Dawson, D.H. and J.A. Nelson. 1977. Wood and bark quality of juvenile jack pine and eastern larch grown under intes?ve culture. Tappi Wood Chem. Conf. For. Biol. Madison, Wi. pp. 221-216. Draper, N.R. and H. Smith. 1981. Applied Regression Analysis. Second Edition. John Wiley and Sons., N.Y. pp. 250-254. Dadswell, H.E. 1958. Wood structure variations occurring during tree growth and their influence on properties. J. Inst. Wood. Sci. (1):11-33. De Guth, E.B. 1980. Relationship between wood density and tree diameter in Pi nus e11iott i i of Misiones, Argentina. Proc. Div. 5. I.U.F.R.O. Meet., Oxford, England. Dinwoodie, J.M. 1961. Tracheid and fiber length in timber. A review of literature. Forestry 34(2):125-144. Duff, G.H. and N.J. Nolan. 1953. Growth and morphogenesis in the Canadian forest species. I. The controls of cambial and apical activity in Pi nus res i nosa Ait. Can. J. Bot. (31):471-513. 80 Elliott, G.K. 1958. Spiral grain in second-growth Douglas-fir and hemlock. Forest Prod. J. 8:205-211. Elliott, G.K. 1970. Wood density in conifers. Commonwealth Forestry Bureau. Oxford, England, Technical Communication No. 8. 44 pp. Erickson, H.D. and T. Arima. 1974. Douglas-fir wood quality studies. Part II. Effects of age and stimulated growth on fibril angle and chemical constituents. Wood Sci. Tech. 8(4):255-265. and A.T. Harrison. 1974. Douglas-fir wood quality studies. Part I. Effects of age and stimulated growth on wood density and anatomy. Wood Sci. Tech. 8(4):207-226. Fielding, J.M. 1967. The influence of si1vicu1tura1 practices on wood properties. Int. Rev. Forestry Res., N.Y. 2:99-126. Foelkel, C.F.B., Barrichelo, L.E.G., Garcia, W. and J.O. Brito. 1976. Kraft cellulose of juvenile and adu11 wood of Pi nus e11ott i i. I.P.E.F., Piracicaba (12):127-142. Forward, D.F. and N.J. Nolan. 1964. Growth and morphogenesis in the Canadian forest species. VII. Progress and control of longitudinal growth of branches in Pi nus res i nosa Ait. Can. J. Bot. 42:923-950. Fowells, H.A. 1965. Si Ivies of Forest Trees of the United States. U.S.D.A. F.S. Agriculture Handbook no. 271. pp. 546-553. Fraser, D.A. 1962. Apical and radial growth of white spruce (Pi cea g1auca (Moench) Voss) at Chalk River, Ontario, Canada. Can. J. Bot. 40:659-668. Freeland, R.O. 1952. Effect of age of leaves upon the rate of photosynthesis in some conifers. Plant. Physiol. 27(4):685-690. Gerhards, CC. 1979. Effect of high-temperature drying on tensile strength of Douglas-fir 2' x 4's. Forest Prod. J. 29(3):39-46. 81 Gerischer, G.F. and A.M. De Villiers. 1963. The effect of heavy pruning on timber properties. For. South Africa. 3:15-41. Goggans, J.F. 1961. The interplay of environment and heredity as factors controlling wood properties in conifers with special emphasis on their effects on specific gravity. Tech. Rept. II. N.C. State Univ., Ra1eigh. 56 pp. Gonzalez, J.S. and R.M. Kellogg. 1978. Evaluating wood specific gravity in a tree improvement program. W.F.P.L. Info. Rep. VP-X.183. Vancouver, B.C. Canada. 14 pp. Hale, J.D. and K.G. Fenson. 1931. The rate of growth and density of the wood of white spruce. Canada Dept. Int. For. Ser. Circ. No. 30. 18 pp. and J.B. Prince. 1940. Density and rate of growth in the spruce and balsam fir of eastern Canada. Canada Dept. Mines and Res. For. Serv. Bul1. 94. 43 pp. Harris, J.M. 1969a. The use of beta rays in determining wood properties. Part 1 - 5. N.Z. J. Sci. 12(2):396-451. . 1969b. On the cause of spiral grain in corewood of radiata pine. N.Z. J. Bot. 7(3):189-213. and H.R. Orman. 1958. The physical and mechanical properties of New Zealand grown Douglas-fir. N.Z. For. Serv. For. Res. Inst. Tech. Pap. No. 24. 87 pp. Haygreen, J.G. and J.L. Bowyer. 1982. Forest Products and Wood Science. The Iowa State University Press. 495 pp. Hudson, D.J. 1966. Fitting segmented curves whose join points have to be estimated. J. Am. Stat. Assoc. 61(316):1097-1129. Husch, B., Miller, C.I. and T.W. Beers. 1982. Forest Mensuration. John Wiley and Sons, Inc., N.Y. pp. 80-89. 82 Ifju G. 1969. Within-growth-ring variation in some physical properties of Southern Pine Wood. Wood Sci. 2(1):11-19. Ifju G. and R.W. Kennedy. 1962. Some variables affecting microtensile strength of Douglas-fir. Forest Prod. J. 12:213-217. , Well wood, G.W. and J.W. Wilson. 1965. Relationship between certain intra-increment physical measurements in Douglas-fir. Tech. Sect. Pulp and Pap. Mag. of Canada 66(9):T475-T483. Isebrands, J.G., Einspahr, D.W., Phelps, J.E. and M.B. Crist. 1982. Kraft pulp and paper properties of juvenile hybrid larch grown under intensive culture. Tappi 65(3):122-126. and CM. Hunt. 1975. Growth and wood properties of rapid-grown Japanese larch. Wood and Fiber 7(2):119-128. Jackson, M. and R.A. Megraw. 1986. Impact of juvenile wood on pulp and paper products. Proc. A Technical Workshop: Juvenile wood - What Does it Mean to Forest Management and Forest Products? F.P.R.S. S.A.F. Bellingham, Wa. pp. 75-81. Jozsa, L.A. and R.M. Kellogg. 1986. An exploratory study of the density and annual ring weight trends in fast-growth coniferous woods in British Columbia. Forintek Canada Corp., Vancouver, B.C. Tech. Rep. 43 pp. and E.P. Swan. 1986. Basic Wood property variation in second-growth Douglas-fir. Extended Abstracts - Second-growth Douglas-fir: Its Utilization and Management for Value. Forintek Canada Corp., Vancouver, B.C. pp. 6-9. Kellogg, R.M. 1982. Coming to grips with wood quality. Forestry Chron. 58(6):254-257. . 1986. A task force approach to our changing resource. Proc. 47349. Managing and Marketing the Changing Resource. F.P.R.S. Conference, Fort Worth, Texas. pp. 64-68. 83 Kellogg, R.M. and R.W. Kennedy. 1986. Implications of Douglas-fir wood quality relative to practical end use. Proc. Symp. on Douglas-fir: Stand Management for the Future. Univ. Wa., Seattle. pp. 97-102. Kennedy, R.W. and J.M. Jaworsky. 1960. Variations in cellulose content of Douglas-fir. Tappi 43(1):25-27. and J.M. Warren. 1969. Within-tree variation in physical and chemical properties of Douglas-fir. Proc. I.U.F.R.O. Second World Consultation of Forest Tree Breeding. F.A.O., Rome. pp. 394-417. Kirk, D.G., Breeman, L.G. and B.J. Zobel. 1972. A pulping evaluation of juvenile loblolly pine. Tappi 55(11):1600-1604. Klinka, K., Nuszdorfer, F.C. and L. Skoda. 1979. Biogeoclimatic units of central and southern Vancouver Island. Province of B.C. M.O.F. I.S.B.N. 0-7719-8213-5. 120 pp. Kozlowski, T.T. 1971. Growth and Development of Trees. Academic Press, N.Y. Vol. I. pp. 1-93, Vol. II. pp. 1-63. Krahmer, R.L. 1966. Variations of specific gravity in western hemlock trees. Tappi 49(5):227-229. . 1986. Fundamental anatomy of juvenile and mature wood. Proc. A Technical Workshop: Junvenile Wood - What does it Mean to Forest Management and Forest Products? F.P.R.S. S.A.F. Bellingham, Wa. pp. 12-16. Larson, P.R. 1962. A biological approach to wood quality. Tappi 45(6):443-448. . 1963. Microscopic wood characteristics and their variations with tree growth. Reprint for work group on wood quality. I.U.F.R.O. Mimeo. 20 pp. . 1964. Some indirect effects of environment on wood formation. In. The Formation of Wood in Forest Trees. M. Zimmermann, Ed. Academic Press, N.Y. pp. 345-365. 84 Larson, P.R. 1965. Stem form of young Lar i x as Influenced by wind and pruning. Forest Sci. 11(4):412-424. . 1969. Wood formation and the concept of wood quality. Yale Univ. Sch. For. Bull. No. 74. 54 pp. . 1973. The physiological basis for wood specific gravity in conifers. Proc. Div. 5. I.U.F.R.O., Stel1enbosch, South Africa. pp. 672-680. Littleford, T.W. 1961. Variation of strength properties within trees and between trees in a stand of rapid-growth Douglas-fir. Forest Prod. Lab. of Can. Rep. V-1028. 20 pp. Loo, J.A., Tauer, C.G. and R.W. McNew. 1985. Genetic variation in the time of transition from juvenile to mature wood in loblolly pine (Pinus taeda L.). Silvae Genetica 34(1):14-19. McKimmy, M.D. 1966. A variation and heretabi1ity study of wood specific gravity in 46-year-old Douglas-fir from known seed sources. Tappi 49(12):542-549. . 1986. The effect of forest management practices on wood properties. Proc. A Technical Workshop: Juvenile Wood - What Does it Mean to Forest Management and Forest Products? F.P.R.S. S.A.F. Bellingham, Wa. pp. 35-49. and R.K. Campbell. 1982. Genetic variation in the wood density and ring width trend in coastal Douglas-fir. Silvae Genetica 31(2-3):43-51. Marts, R.W. 1949. Effect of crown reduction on taper and density in long leaf pine. South Lumberman 179:206-209. Megraw, R.A. 1985. Wood Quality Factors in Loblolly Pine - The Influence of Tree Age, Position in Tree and Cultural practice on Wood Specific Gravity, Fiber Length and Fibril Angle. Tappi Press, Atlanta, Ga. 88 pp. . 1986. Douglas-fir wood properties. Proc. Symposium on Douglas-fir Stand Management for the Future. Univ. Wa., Seattle. pp.81-95. 85 Megraw, R.A. and W.T. Nearn. 1972. Detailed D.B.H. density profile of several trees from Douglas-fir fertilizer/thinning plots. Proc. Symp. on Effect of Growth Acceleration on Properties of Wood. U.S.D.A. Forest Prod. Lab., Madison, Wi. 12 pp. Meylan, B.A. 1968. Cause of high longitudinal shrinkage in wood. Forest Prod. J. 18(4):75-78. Mitchell, K.J. 1969. Simulation of the growth of even-aged stands of white spruce. Yale Univ. Sch. For. Bull. No. 7. 47 pp. . 1975. Dynamics and simulated yield of Douglas-fir. Forest Sci. Mono. 17. 39 pp. . 1980. Distance dependent individual tree stand models: Concepts and applications. Proc. Workshop in Forecasting Forest Stand Dynamics. School of Forestry, Lakehead Univ., Thunder Bay, Ont. pp. 100-137. and I.R. Cameron. 1985. Managed stand yield tables for coastal Douglas-fir: Initial density and precommercial thinning. B.C. Ministry of Forests Research Branch. Land Management Rep. ISSN 0702-9861. No. 31. 69 pp. Moody, R.C. 1970. Tensile strength of finger joints in pith-associated and non-pith-associated southern pine lumber. U.S.D.A. For. Ser. Res. Pap. F.P.L. 138. Forest Prod. Lab., Madison, Wis. 11 pp. Northcott, P.L. 1957. Is spiral grain the normal growth pattern? Forestry Chron. 33(4):335-352. Oliver, CD. 1986. Silviculture and juvenile wood. Proc. A Technical Workshop: Juvenile Wood - What Does it Mean to Forest Management and Forest Products? F.P.R.S. S.A.F. Bellingham, Wa. pp. 29-34. Owens, J.N. 1968. Initiation and development of leaves in Douglas-fir. Can. J. Bot. 46:271-278. Panshin, A.J. and C. de Zeeuw. 1980. Textbook of Wood Technology. Fourth Edition. McGraw-Hill Book Co., N.Y. pp. 201-240. 86 Parker, M.L. and L.A. Jozsa. 1973. X-ray scanning machine for tree-ring and density analysis. Wood and Fiber 5(3):192-197. Parker, M.L., Schoor1emmer, J. and L.J. Carver. 1973. A computerized scanning densitometer for automatic recording of tree-ring width and density data from x-ray negatives. Wood and Fiber 5(3):237-248. Paul, B.H. 1950. Wood quality in relation to site quality of second-growth Douglas-fir. J. Forestry 48(3):175-179. . 1957. Juvenile wood in conifers. U.S.D.A. Forest Prod. Lab. Rep. 2094. 6 pp. . 1960. The juvenile core in conifers. Tappi 43(1):1-2. Pearson, E.S. and H.O. Hartley. 1954. Biometrika. Tables for Statisticians, Vol. 1. Cambridge Univ. Press, N.Y. pp. 290-299. Pearson, R.G. and R.C. Gil more. 1971. Characteristics of the strength of juvenile wood of loblolly pine (Pinus taeda L.). Forest Prod. J. 21(l):23-30. . 1980. Effect of fast-growth rate on the mechanical properties of loblolly pine. Forest Prod. J. 30(5):47-54. and B.E. Ross. 1984. Growth rate and bending properties of selected loblolly pines. Wood and Fiber Sci. 16(l):34-47. Perry, T.O. and CW. Wang. 1958. Variation in the specific gravity of slash pinewood and its genetic and silvicultural implications. Tappi 41(4):178-180. Plumptre, R.A. and S. Austin. 1978. The influence of pruning on the density and rate of growth of Pi nus  patu1 a. Proc. I.U.F.R.O. Conf. Wood Quality and Tropical Species. F.W. Tamolang, Ed. pp. 351-370. Polge, H. 1964. The juvenile wood of conifers. Rev. For. Franc. 16:474-505. 87 Polge, H., Keller, R. and F. Thiercelin. 1973. Influence de l'elagage de branches vivantes sur la structure des aeroissements annuels et sur quelques characteristiques du bois de douglas et de grandis. Ann. Sci. For. 30:127-140. Reeb, D. 1984. Influence of spacing and artificial pruning on the production of clearwood of Douglas-fir. Unpub. M.F. Thesis. Dept. of Forestry, Univ. B.C. 255 pp. Rendle, B.J. 1958. A note on juvenile and adult wood. I.A.W.A. News Bull. 2:1-6. . 1959. Fast-grown coniferous timber - some anatomical considerations. Quart. J. Forestry 53:116-122. . 1960. Juvenile and adult wood. J. Inst, of Wood Sci. 5:58-61. and E.W. Phillips. 1957. The effect of rate of growth (ring width) on the density of softwoods. Forestry 31:113-120. Reukema, D.L. 1959. Missing annual rings in branches of young growth Douglas-fir. Ecol. 40:480-482. . 1961. Crown development and its effect on stem growth of six Douglas-fir. J. Forestry 59(5):370-371. Richardson, S.D. 1961. A biological basis for sampling in studies of wood properties. Tappi 44(3):170-173. Richardson, S.D. and J.M. Dinwoodie. 1960. Studies on the physiology of xylem development. I. The effect of night temperature on tracheid size and wood density in conifers. J. Int. Wood. Sci. 6:3-13. Rutter, A.J. 1957. Studies in the growth of young plants of Pinus sy1vestr i s L. I. The annual cycle of assimilation and growth. Ann. Bot. 21(83):399-426. Sastry, CB. and R.W. Well wood. 1971. Individual tracheid weight-1ength relationships in Douglas-fir. Tappi 54(10):1686-1690. 88 Schmidt, J.D. and W.J. Smith. 1961. Wood quality evaluation and improvement in Pinus caribaea Morelet. Queensland For. Serv. Res. Note No. 15. 59 pp. Senft, J.F., Bendtsen, B.A. and W.L. Galligan. 1985. Weak wood, fast-grown trees make problem lumber. J. Forestry 83(8):476-484. , Quanci, M.J. and B.A. Bendtsen. 1986. Property profile of 60-yeai—old Douglas-fir. Proc. A Technical Workshop: Juvenile wood - What Does it Mean to Forest Management and Forest Products? F.P.R.S. S.A.F. Bellingham, Wa. pp. 17-28. Shiokura, T. 1984. The classification of Juvenile wood and its perimeter in coniferous trees. Proc. Pacific Reg. Wood Anat. Conf. Tsukuba, Ibariki, Japan. pp. 76-78. Smith, D.M. 1954. Maximum moisture content method for determining specific gravity of small wood samples. U.S.D.A. For. Serv., Forest Prod. Lab. Rep. 2014. 8 pp. . 1968. Wood quality of loblolly pine after thinning. U.S.D.A. Forest Prod. Lab. Madison, Wi. Res. Pap. F.P.L. 89. 10 pp. Smith, J.H.G., Heger, L. and J. Hejjas. 1966. Patterns in growth of earlywood, latewood and percentage latewood determined by complete analysis of 18 Douglas-fir trees. Can. J. Botany. 44:453-466. , Ker, J.W. and Csizmazia, J. 1961. Economics of reforestation of Douglas-fir, western hemlock and western redcedar in the Vancouver forest district. Fac. of Forestry, Univ. of B.C. For. Bulletin No. 3. 144 pp. Smith, R.W. and D.G. Briggs. 1986. Juvenile wood: Has it come of age? Proc. A Technical Workshop: Juvenile Wood - What Does it Mean to Forest Management and Forest Products? F.P.R.S. S.A.F. Bellingham, Wa. pp. 5-9. Taylor, F.W., Wang, E.I.C., Yanchuk, A. and M.M. Micko. 1982. Specific gravity and tracheid length variations of white spruce in Alberta. Can. J. Forestry Res. 12:561-566. 89 Trendelenberg, R. 1935. Variations in the density of important coniferous timbers due to locality, habitat, and difference in individual trees. Zeitschrift des Vereines Deutscher Ingenieure. Imp. For. Inst. Univ. Oxford. Translation No. 1. 79(4):85-89. Turnbull, J.M. 1937. Variations in strength of pine timbers. S. African J. Sci. 33:653-682. Wang, E.I.C. and M.M. Micko. 1984. Wood quality of white spruce from north-central Alberta. Can. J. Forestry Res. 14:181-185. Wangaard, F.F. and F.V. Zumwalt. 1949. Some strength properties of second-growth Douglas-fir. J. Forestry 47(1):18-24. Wareing, P.F. 1958. The physiology of cambial activity. J. Inst. Wood Sci. 1:34-42. Wei 1 wood, R.W. 1952. The effect of several variables on the specific gravity of second-growth Douglas-fir. Forestry Chron. 28(2):34-42. . 1960. Specific gravity and tracheid length variations in second-growth western hemlock. J. Forestry 58(5):361-368. and P.E. Jurazs. 1968. Variation in sapwood thickness, specific gravity and tracheid length in western redcedar. Forest Prod. J. 18(12):37-46. and J.H.G. Smith. 1962. Variation in some important qualities of wood from young Douglas-fir and hemlock trees. Fac. of For., Univ. Brit. Col., Vancouver. Res. Pap. No 50. 15 pp. Wilcox, H. 1962. Cambial growth characteristics. J_n Tree Growth. T.T. Kozlowski, Ed. Ronald Press, N.Y. pp. 57-88. Wilson, B.F. 1984. The Growing Tree. The University of Massachusetts Press. 138 pp. Wodzicki, T. 1960. Investigation on the kind of Lar i x  po1 onica Rac. wood formed under various photoperiodic conditions. I. Plants growing under natural conditions. Acta. Soc. Bot. Poloniae. 29:713-730. 90 Wood, F.F. and J. Bryan. 1960. The silviculture and quality of Sitka spruce grown in Great Britain. Proc. Fifth World Forest. Cong. Univ. Wa. Seattle. Vol 111:1372-1374. Woodfin, R.O. Jr. 1969. Spiral grain patterns in coastal Douglas-fir. Forest Prod. J. 19(l):53-60. Yang, K.C., Benson, CA. and J.K. Wong. 1986. Distribution of juvenile wood in two stems of Lar i x 1ar i c i na. Can. J. Forestry Res. 16:1041-1049. Zobel, B.J. 1984. The changing quality of the world wood supply. Wood Sci. Techno 1. 18:1-17. and R.C Kellison. 1972. Short-rotation forestry in the southeast. Tappi 55(8):1205-1208. and D.G. Kirk. 1972. Proc. Symp. on the Effect of Growth Acceleration on the Properties of Wood. U.S.D.A. F.P.L. Madison, Wi. pp. M1-M20. and R.L. McElwee. 1958. Natural variation in wood specific gravity of loblolly pine and an analysis of contributing factors. Tappi 41(4):158-161. , Matthias, M., Roberds, J.H. and R.C. Kellison. 1968. Moisture content of southern pine trees. School For. Res. N.C State Univ. Raleigh, N.C Tech. Rep. No. 37. 44 pp. and J.T. Talbert. 1984. Applied Forest Tree Improvement. John Wiley and Sons, Inc. pp. 376-413. , Webb, C. and F. Henson. 1959. Core or juvenile wood of loblolly and slash pine trees. Tappi 42(5):345-356. 91 TabIt 1. Douglas-fir stand characteristics. Location Coordinates Biogeo- Organization Ag* Height Crown Crow 6.B.H. Density Basal Douglas-Latitude Longitude cliaatic Sad. Length (stei/ha) Area fir Zone (B.H.) (•) (•) (•) (cn) ha) (• sq/ Z ha) (1) (2) (2) (2) (2) (2) (3) (3) (3) o o Ladysnith 49 OO'H 123 52'W CDFb Crovn Forest 54 42.2 S.7 17.3 54.5 Industries o o Cassidy 49 04'N 123 55'y CDFb MacMillan 58 41.7 4.6 17.7 57.8 (54 46.1 100 Bloedel o o Caipbell 50 OO'N 125 17'W CDFb B.C. Forest 53 38.8 4.2 17.5 52.7 395 43.3 100 River Products o o Lake 48 48'N 124 09'W CWHal C.I.P. Inc. 39 33.6 3.4 17.1 38.4 381 42.1 43 Covichan o o Lizard 48 35'N 124 08'W CWHbl B.C. Forest 46 35.6 3.3 18.1 45.8 705 50.7 34 Late Products o o Jordan 48 24>N 124 08'W CWHbl - Western Forest 52 40.3 3.4 15.9 52.5 523 51.8 40 River CWHal Products o o Haney 49 17'N 122 30'W CWHa2 - U.B.C. Research 48 7CWHb2 Forest o o Hart 50 02'N 125 20'H CDFb B.C. Ministry 37 100 of Forests o o Henekay 50 02'N 125 19'W CDFb B.C. Ministry 35 10of Forests (1) Biogeoclinatic Zone Classification (Klinka et al., 1979). CDFb: Coastal Douglas-fir - vetter subzone CWHal: Coastal Western Henlock - Vancouver Island, wetter subzone CWHa2: Coastal Western Henlock - Pacific Range, vetter subzone CWHbl: Coastal Western Henlock - Windvard Subnontane Naritine, drier subzone CWHb2: Coastal Western Henlock - Windward Montana Haritine, drier subzone (2) Based on the average of ten dominant and codoninant trees per stand. (3) Based on prist cruise information. Table 2. Saaple tree characteristics. Sample Location Age D.B.H. Height Collection Contents Tree Mo. (B.H.) Bate (ca) (a) IAS Ladysaith 56 46.8 37.0 Nay, 19B5 fecond-grovth IA7 Ladysaith SI 46.1 37.0 Nay, 1985 second-growth IK Cassidy 53 49.2 44.7 Hay, 1983 second-growth IBM Cassidy 63 49.1 36.1 Hay, 1985 second-grovth 1C1 Caapbell River 54 47.3 40.0 Hay, 1985 second-grovth IC6 Caapbell River 53 49.1 38.0 Hay, 1985 SKond-growth 1D5 Lake Covichan 33 45.5 32.0 June, 1985 second-grovth IK lake Covicha* 35 43.2 29.5 Jane, 1985 second-growth 1E3 Lizard Lake 43 39.4 35.0 June, 1985 second-grovth IE8 Lizard Lake 47 68.2 36.0 Jane, 1985 second-growth 1F7 Jordan River 53 57.5 41.5 June, 1985 second-grovth trio Jordan River 52 52.7 39.0 Jane, 1985 second-grovth RFl Haney 21 26.8 23.3 April, 1985 SKond-grovth RF2 Haney 51 68.3 31.7 April, 1985 second-growth pruned RF3 Haney 45 44.0 29.7 January, 1986 second-grovth pruned CRl Hart 37 19.8 16.8 October, 1985 plantation-grown CR2 Heaekay 35 36.7 31.5 October, 1985 plantation-grown 93 Table 3. Tree crovn characteristics. Saaple Location Height Age Crovn Crovn Base Crovn Average Average Average Crovn Coefficient Tree No. (B.H.) Length Height Base Crovn Crovn Crovn Radius Values Age Length Height Age (BL) (a) (a) (a) (L) (a) (a) (b) (a) (1) (1) (1) (2) (2) (2) IAS Ladysaith 37.0 56 14.8 22.2 28 14.3 22.7 27 4.65 3.952 1A7 Ladysaith 37.0 51 13.6 23.4 29 10.0 27.0 23 4.68 4.956 1B6 Cassidy 44.7 53 21.9 22.8 28 13.0 31.7 17 4.23 3.806 IBil Cassidy 36.1 63 15.3 20.8 35 15.1 21.3 34 4.40 3.625 1CI Caapbell River 40.0 54 15.9 24.1 29 13.8 26.4 26 3.61 3.612 1C6 Caapbell River 30.0 53 13.2 24.8 24 11.1 26.9 19 3.70 3.701 IBS Lake Covichan 32.0 33 16.9 IS.l 20 12.0 19.9 13 3.47 3.262 IK Lake Covichan 29.5 35 11.3 18.2 16 8.5 21.0 13 3.40 3.966 1E3 Lizard Lake 35.0 43 17.9 17.1 21 13.2 22.2 17 4.01 3.588 1E8 Lizard Lake 36.0 47 14.4 21.6 25 8.8 27.2 17 3.66 4.204 1F7 Jordan River 41.5 53 18.1 23.4 28 7.7 33.8 15 3.24 4.080 1F10 Jordan River 39.0 52 14.2 24. B 29 8.7 30.3 21 2.91 3.362 Rf 1 Haney 23.3 21 14.3 9.0 12 8.1 15.2 6 2.64 3.213 RF2 Haney 31.7 51 21.0 10.7 39 - - - - -RF3 Haney 29.7 45 18.1 11.6 33 - - - - -CR1 Hart 16.8 37 6.7 9.4 22 3.9 12.8 16 1.4B 3.239 CR2 Heaekay 31.5 35 12.8 18.7 19 5.6 25.9 9 2.80 4.420 Averages (3) 34.5 46 14.7 19.7 24 10.2 24.2 18 3.50 3.799 (1) Base of the live crovn height observed as the lovest part of the crovn with three or tore live branches. (2) Average crovn height estiaated as tree height ainus L, where L is given by the following equation: BL / (bd) L = c ( e -1) L ' crown length or distance froa terainal leader to branch base c e coefficient describing the shape of the crovn (estiaated for Douglas-fir as (.1 a) BL - crown radiis b - coefficient relating branch growth to height growth d 8 coefficient coapensating for branch crooks (estiaated for Douglas-fir as 0.975) e =2.71828 (3) Overall averages not including pruned trees RF2 and RF3. 94 Table 4. Sunnary of relative density values for sections at breast height (B.H.). Satple Location Height Nunber of Relative Density Tree Ho. 6rowth (•) Increaents Mean S.D. Haxiaua Hiniaua IAS Ladysnith 1.30 56 .542 .062 .640 .380 1A7 Ladysnith 1.30 51 .563 .057 .660 .430 IBS Cassidy 1.30 53 .520 .055 .650 .420 1BU Cassidy 1.30 63 .543 .057 .680 .430 1C1 Catpbeil River 1.30 54 .558 .053 .660 .440 1C6 Catpbell River 1.30 53 .566 .062 .700 .440 IDS Lake Covicban 1.30 33 .474 .034 .540 .420 1D6 Lake Covichan 1.30 35 .488 .062 .600 .380 1E3 Lizard Lake 1.30 43 .595 .049 .680 .500 1E8 Lizard Lake 1.30 47 .487 .044 .590 .390 1F7 Jordan River 1.30 53 .551 .061 .650 .430 1F10 Jordan River 1.30 52 .537 .060 .640 .430 RF1 Haney 1.30 21 .429 .041 .530 .360 CR1 Hart 1.30 37 .536 .070 .670 .410 CR2 Henekay 1.30 35 .444 .039 .510 .340 All Trees 1.30 46 .523 .054 .625 .413 95 Table 5. Suaaary of relative density values for sections saapled at 20 percent of total height. Saaple Tree No. Location Height (a) Nuaber of firovth Increments Mean Relative Density S.D. Haxiaua Hiniaua 1A5 Ladysaith 7.55 47 .468 .063 .620 .360 1A7 Ladysaith 7.20 44 .520 .071 .650 .370 1B6 Cassidy 8.55 45 .504 .055 .590 .400 1BU Cassidy 7.10 53 .489 .037 .590 .410 1C1 Caapbell River 8.00 47 .494 .049 .570 .390 1C6 Caapbell River 7.60 46 .502 .059 .600 .380 1D5 Lake Covichan 6.40 27 .482 .039 .560 .430 1D6 Lake Covichan 5.90 29 .437 .043 .520 .350 1E3 Lizard Lake 7.10 32 .498 .03B .590 .440 1E8 Lizard Lake 7.20 41 .463 .057 .610 .360 1F7 Jordan River 8.30 46 .490 .061 .610 .380 IF 10 Jordan River 7.80 46 .485 .052 .570 .390 RF1 Haney 4.54 17 .423 .035 .500 .380 Cftl Hart 3.34 34 .543 .076 .680 .420 CR2 Heaekay 6.40 29 .483 .049 .580 .370 All Trees 6.85 39 .485 .052 .589 .389 96 Table 6. Suaaary of relative density values for sections saapled at 40 percent of total height. Saaple Location Height Nuaber of Relative Density Tree Ho. Srovth (a) Increments Hean S.D. Haxiaua Hiniaua IAS Ladysaith 14.90 40 .445 .049 .540 .370 IA7 Ladysaith IS. 10 39 .480 .045 .570 .380 IBS Cassidy 17.20 32 .495 .042 .570 .420 mi Cassidy 14.20 45 .478 .033 .580 .410 1C1 Caapbell River 16.00 39 .468 .032 .520 .400 1C6 Caapbell River 15.20 36 .496 .040 .560 .430 IDS Lake Covichan 12.80 20 .475 .035 .550 .430 ID6 Lake Covichan 11.90 23 .450 .037 .550 .400 1E3 Lizard Lake 14.00 26 .450 .024 .510 .390 IE8 Lizard Lake 14.40 33 .456 .039 .530 .370 1F7 Jordan River 16.40 38 .473 .040 .570 .420 IF10 Jordan River 15.90 39 .472 .030 .530 .410 RF1 Haney 9.00 13 .398 .054 .500 .340 Cfil Hart S.I0 31 .486 .069 .620 .390 CR2 Heaekay 25.77 22 .488 .028 .540 .440 All Trees 13.59 32 .467 .040 .549 .400 97 Tab 1 e 7. Sumry of relative density values for sections sampled at 60 percent of total height. Saaple Tree No. Location Height (a) Nuaber of 6rovth Increments Hean Relative Density S.D. Haximua Hinimum 1A5 Ladysaith 22.2 28 .460 .038 .560 .400 1A7 Ladysaith 22. S 31 .479 .054 .670 .410 IB6 Cassidy 25.8 24 .478 .027 .540 .440 1611 Cassidy 21.3 29 .479 .031 .560 .430 1C1 Caapbell River 24.1 29 .448 .023 .510 .410 IC6 Caapbell River 22.8 25 .478 .034 .550 .410 IDS Lake Covichan 19.2 14 .456 .034 .510 .390 106 Lake Covichan 17.7 17 .428 .041 .530 .360 1E3 Lizard Lake 21.0 17 .417 .017 .450 .400 IE8 Lizard Lake 21.6 25 .420 .034 .500 .360 1F7 Jordan River 24.9 22 .497 .056 .650 .430 IF 10 Jordan River 23.4 26 .495 .026 .550 .460 RF1 Haney 13.6 8 .445 .042 .510 .400 CRl Hart 9.9 17 .485 .025 .540 .450 CR2 Heaekay 24.8 20 .459 .048 .630 .400 All Trees 20.6 22 .462 .035 .551 .410 98 Table 8. Suttary of relative density values for sections sanpled at 80 percent of total height. Saaple Tree No. Location Height (n) Nunber of Grovth Increments Hean Relative Density S.D. Haxinun Hininun IAS Ladysnith 29.4 13 .440 .030 .510 .410 IA7 Ladysnith 29.9 18 .499 .038 .600 .450 IK Cassidy 36.1 12 .479 .052 .610 .420 IBM Cassidy 28.4 19 .488 .032 .570 .450 1C1 Canpbell River 32.0 15 .411 .015 .460 .400 1C6 Canpbell River 30.4 12 .499 .035 .560 .430 IDS Lake Covichan 25.6 8 .540 .027 .580 .500 1D6 Lake Covichan 23.6 8 .468 .027 .510 .420 1E3 Lizard Lake 28.0 9 .483 .031 .560 .450 1E8 Lizard Lake 28.8 14 .454 .030 .530 .420 1F7 Jordan River 33.7 14 .517 .058 .640 .450 If 10 Jordan River 31.2 19 .420 .042 .560 .430 Rf 1 Haney 18.2 4 .492 .025 .520 .460 CR1 Hart 13.4 7 .485 .040 .540 .430 CR2 Henekay 25.8 16 .493 .041 .590 .440 All Trees 27.6 13 .478 .035 .556 .437 99 Table 9. Sunaary of the distribution of relative density profiles which showed no juvenile - nature wood transition. Section Ranter «f Height Juvenile Hood 6rovtn Increment Height Sections Hean S.I. Hai. Hin. Hean S.I. (tax. Hin. B.H. 1 1.30 - 1.30 1.30 21.00 - 21.00 21.00 201 1 4.54 - 4.54 4.54 17.00 - 17.00 17.00 401 6 10.83 3.31 14.40 5.10 23.67 7.37 33.00 13.00 602 15 20.59 4.26 25.80 9.95 22.13 6.50 31.00 8.00 801 IS 27.63 5.83 36.10 13.38 12.53 4.58 19.00 4.00 All Sections 38 20.99 8.58 36.10 1.30 18.42 7.45 33.00 4.00 100 Table 10. Transition determination by coaparing simple against segmented linear regression aodels on relative density profiles. Sample Location Section Juvenile - Simple Linear Hodel 1 Segmented Linear Hodel 2 'F* Value •F" Prob. Tree Height Nature Hood No. Transition Hean Square Hean Square Age -3 -3 Hodel 1 / (years) D.F. ( i 10 ) D.F. ( x 10 ) Hodel 2 P > F 1A5 Ladysnith B.H. 22 50 2.081 48 1.623 1.281 0.20 201 28 41 1.971 39 1.857 1.061 0.42 401 25 23 1.660 21 1.480 1.121 0.40 1A7 Ladysnith B.H. 19 41 2.058 39 1.468 1.401 0.14 20Z 24 35 2.609 33 1.743 1.496 0.14 401 24 31 1.303 29 0.841 1.548 0.10 m Cassidy B.H. 16 44 1.302 42 1.129 1.153 0.32 201 17 35 0.751 33 0.666 1.128 0.37 40Z 18 24 0.564 22 0.542 1.055 0.40 1 BI 1 Cassidy B.H. 32 56 1.738 54 1.247 1.394 0.11 201 25 50 1.373 48 0.863 1.590 0.05 40Z 16 41 1.176 39 1.023 1.150 0.34 1C1 Campbell B.H. 19 47 1.182 45 0.915 1.292 0.19 River 20Z 25 41 0.868 39 0.597 1.454 0.02 40Z 29 32 0.482 30 0.463 1.042 0.42 IC6 Campbell B.H. 15 47 2.258 45 1.980 1.141 0.33 River 20Z 35 41 2.247 39 1.813 1.239 0.26 401 26 31 2.284 29 1.006 2.270 0.00 1D5 Lake B.H. 19 30 0.861 28 0.844 1.020 0.43 Covichan 20Z 20 23 0.759 21 0.712 1.070 0.44 101 Table 10. Transition determination by comparing simple against segaented linear regression models on relative density profiles (cont.). Saaple Location Section Juvenile - Simple Linear Model 1 Segaented Linear Model 2 V Value •r Prob. Tree Height Nature Hood No. Transition Mean Square Mean Square Age -3 -3 Model 1 / P > F (years) D.F. ( x 10 ) D.F. ( x 10 ) Model 2 106 Lake B.H. 17 30 1.015 28 0.878 1.156 0.35 Covichan 20Z IS 24 0.661 22 0.542 1.220 0.31 1E3 Lizard B.H. 28 32 1.069 30 0.985 1.085 0.41 Lake 20Z 26 25 0.785 23 0.762 1.031 0.47 401 11 19 0.405 17 0.293 1.378 0.25 IE8 Lizard B.H. 34 41 I.0S5 39 0.930 1.134 0.35 Lake 20Z 26 31 1.220 29 1.019 1.197 0.32 1F7 Jordan B.H. 25 45 1.633 43 0.801 2.037 0.01 River 201 25 39 1.086 37 0.422 2.573 0.00 40Z 24 28 1.418 26 0.852 1.664 0.10 1F10 Jordan B.H. 24 49 1.338 47 0.985 1.358 0.15 River 20Z 22 39 1.465 37 1.023 1.425 0.14 401 15 35 0.557 33 0.448 1.244 0.27 CXI Hart B.H. 20 28 1.576 26 1.331 1.183 0.2S 20Z IS 22 1.905 20 1.019 1.873 0.07 CR2 Heaekay B.H. 23 24 0.930 22 0.797 1.166 0.32 20Z 13 23 0.866 21 0.798 1.087 0.42 102 TabI* 11. Sutury of tht distribution of relative density profiles which shoved juvenile - nature vood transition. Section Nunber of Height Total Nunber No. of Juvenile-Nature Mood Height Sections (•) of 6rovth Increments Trans. Growth Increments Hean S.D. Hai. Him. Mean S.D. Has. Hin. Hean S.D. Hax. Hin. B.H. 14 1.30 - 1.30 1.30 47.50 9.33 63.00 33.00 22.36 5.77 34.00 15.00 201 14 7.03 1.03 8.55 3.34 40.43 8.44 53.00 27.00 22.86 5.72 35.00 13.00 401 9 15.43 1.04 17.20 14.00 37.11 5.40 45.00 26.00 20.89 6.05 29.00 11.00 All Sections 37 6.91 5.59 17.20 1.30 42.30 9.07 63.00 26.00 22.18 5.71 35.00 11.00 103 Table 12. Suuary of the relative density values of the profiles which shoved no juvenile - nature wood transition. Section Height Nuaber of Sections Nuaber of Growth Increaents Relative tensity Juvenile Hood Nature Hood Total Juvenile Hood Hatare Wood Total Hean S.B. Hax. Hin. Hean S.B. Hax. Hin. Heaa S.B. Hax. Hin. B.H. 1 21.00 - 21.00 .429 .041 .530 .360 - .429 .041 .530 .360 201 1 17.00 - 17.00 .423 .035 .500 .380 - - - .423 .035 .500 .380 401 6 23.66 - 23.66 .459 .044 .548 .39S - .459 .044 .548 .395 60Z 15 22.13 - 22.13 .462 .035 .551 .410 - .462 .035 .551 .410 801 15 18.42 - 18.42 .481 .035 .556 .437 - .481 .03S .556 .437 All Sections i 38 18.42 18.42 .467 .031 .551 .416 .467 .031 .551 .416 104 Table 13. Suaaary of the relative density values of the profiles which showed juvenile - aature wood transition. Section Huaber of Average Hunter of Relative tensity Height Sections Growth Increaents Juvenile Hood Hature Hood Total Juvenile Hature Total Hean S.B. Max. Hin. Mean S.D. Max. Win. Mean S.D. Max. Min. Hood Hood I.H. 14 22.36 24.71 47.30 .491 .046 .601 .417 .563 .034 .630 .496 .529 .055 .634 .417 201 14 22.S6 17.57 40.43 .460 .046 .566 .389 .529 .032 .592 .474 .490 .054 .596 .389 401 9 20.89 16.22 37.11 .461 .037 .537 .406 .491 .027 .540 .439 .473 .037 .550 .403 All Sections 37 22.18 19.95 42.30 .472 .044 .572 .404 .533 .032 .594 .474 .501 .050 .599 .403 105 TABLE 14. Overall sunnary of relative density values for juvenile and nature wood. Section Nunber of Average Nunber of Relative Oensity ght Sections Growth Increaents Juvenile Hood Nature Hood Total Juvenile Hood Hatare Hood Total Hean S.D. Hai. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. B.H. IS 22.67 24.71 45.73 .487 .046 .596 .413 .563 .034 .630 .496 .523 .054 .625 .413 201 15 22.47 17.57 38*86 .457 .045 .561 .389 .529 .032 .592 .474 .465 .052 .589 .389 401 IS 22.00 16.22 31.73 .460 .040 .541 .401 .491 .027 .540 .439 .467 .040 .549 .400 601 IS 22.13 - 22.13 .462 .035 .551 .410 - .462 .035 .551 .410 801 IS 12.S3 12.53 .481 .035 .556 .437 .... .481 .035 .556 .437 All Sections 75 20.28 19.95 30.20 .469 .040 .561 .410 .533 .032 .594 .474 .483 .043 .547 .410 106 Table 15. Suaaary of sampled dead branch characteristics. Tret Number of Height Position Number of Number Branches (m) 6rovth Increments Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. IAS 5 17.8 4.218 22.2 12.2 13.4 5.983 20.0 6.0 1A7 7 (S.S 7.378 23.4 5.2 11.1 5.550 22.0 6.0 1B6 11 13.0 (.374 22.0 3.4 12.0 2.932 18.0 B.O 1811 13 11.2 ..037 20.8 2.4 18.3 S.360 26.0 11.0 1C1 10 17.4 5.316 24.1 8.0 16.5 4.648 24.0 10.0 IC6 11 16.6 4.576 24.1 10.5 14.2 4.051 22.0 9.0 IDS 9 8.3 4.431 1S.1 2.0 10.7 1.481 13.0 9.0 1D6 9 10.0 5.394 17.7 2.7 10.5 2.006 15.0 8.0 1E3 10 8.4 4.773 16.0 2.2 10.2 2.699 15.0 7.0 1E8 12 12.9 S.886 21.6 3.3 14.6 2.839 19.0 10.0 1F7 12 12.4 6.640 22.4 2.5 11.0 3.604 18.0 7.0 IF 10 11 13.9 6.311 23.4 4.2 8.4 2.383 13.0 6.0 All Trees 120 12.9 6.226 24.1 2.0 12.7 4.628 26.0 6.0 107 libit 16. Suaaary of lovest relative density values. Tree Hunter of Height Position Lovest Relative Density Nutber Sections (•) Number of 6rovth Increments Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1A5 S 1S.1 1A7 5 1S.2 1B6 5 17.8 1111 5 14.S 1C1 5 16.3 1C6 5 IS.S 1D5 S 13.1 1D6 5 12.1 1E3 5 14.3 1E8 3 14.7 1F7 S 16.9 1F10 S 1S.9 CR1 S 6.6 CR2 5 12.8 11.207 29.4 1.3 11.474 29.5 1.3 13.757 36.1 1.3 10.839 28.3 1.3 12.269 32.0 1.3 11.613 30.4 1.3 9.717 25.6 1.3 8.926 23.6 1.3 10.648 28.0 1.3 10.981 28.8 1.3 12.882 33.7 1.3 11.928 31.2 1.3 4.954 13.4 1.3 9.714 25.8 1.3 7.4 2.302 11.0 5.0 7.2 1.483 9.0 5.0 6.8 2.168 9.0 4.0 6.0 3.336 10.0 2.0 5.2 0.837 6.0 4.0 5.8 2.049 8.0 4.0 5.8 4.438 13.0 2.0 7.2 4.087 13.0 4.0 7.0 1.871 10.0 5.0 9.8 3.834 15.0 5.0 7.4 1.140 9.0 6.0 7.4 5.367 14.0 2.0 9.0 1.581 11.0 7.0 7.8 1.924 10.0 5.0 All Trees 70 14.3 10.229 36.1 1.3 7.1 2.904 15.0 2.0 108 Table 17. Differences between height of juvenile - nature wood transition point, height of crovn base and total height. Section Height Difference Between Transition Height Difference Between Transition Height Point and Crovn Base Point and'Total Height I Obs. Hean S.D. Hax. Hin. I Obs. Hean S.D. Hax. Hin. B.H. 12 10.25 3.802 17.22 8.09 14 18.95 5.875 28.97 8.65 20Z 12 8.71 3.404 13.23 2.52 14 18.51 4.324 23.54 7.23 401 9 3.85 3.280 6.72 -3.15 9 14.59 4.081 22.02 9.78 All Sections 33 7.95 4.311 17.22 -3.15 37 17.72 5.118 28.97 7.23 109 Tib It 18. Suaaary of height difference statistics. first Height Second Height Nutter Height Difference (•) Heasureaent Heasuretent of Observations Hean S.B. Hax. Hin. Total height Live crovn base height IS 14.78 3.399 21.90 6.80 Total height Average crovn height IS 10.26 3.295 15.10 4.00 Average crovn height Live crovn base height 15 4.50 2.920 10.40 0.20 Total height Average transition A 14 12.63 3.835 18.76 7.13 Total height Average transition B 14 17.91 4.172 25.28 8.06 Average transition A Average transition B 14 5.28 4.388 16.35 0.11 Average transition A Live crovn base height 14 2.16 3.658 8.90 -4.36 Average transition A Average crovn height 14 -2.21 3.971 -9.96 5.17 Average transition B Live crovn base height 14 -3.12 3.737 -10.88 3.04 Average transition B Average crovn height 14 -7.50 4.993 -17.94 -1.52 110 Table 19. Correlation natrix for live crovn base height, average crovn height, average transition height A and average transition height B. ! Live Crovn ! Average ! Average ! Average ! Base Height ! Crovn Height ! Transition • Transition I 1 ! Height A 1 Height B Live Crovn Base Height 1.0000 Average Crovn Height 0.8402 It 1.0000 Average Transition Height A 0.8419 II 0.7936 II 1.0000 Average Transition Height B 0.7340 tl 0.5888 1 0.7407 II 1.0000 I Significant at the 0.05 level II Significant at the 0.01 level (14 observations) Ill Figure 1. Location of Douglas-fir stands. 112 o.e f i i i i i i ; i i i i 5 10 IS 20 25 SO 35 40 45 50 55 GROWTH INCREMENTS FROM PITH Figure 2. Pith to bark relative density profiles for sanple tree IAS. 113 1MEIGMT»B.H7| -1 1 : 1 1 1 1 I 1 1 1— 5 10 15 20 25 30 35 40 45 50 55 GROWTH INCREMENTS FROM PITH Figure 3a. Segmented linear regression aodel or. pith to bark relative density profile for a sample radius. Figure 3b. Simple linear regression model on pith to bark relative density profile for a saaple radius. •t •It 4? <0t tt 1M It HI 1J IK JO lit >tt M lit »' Itt tt its Tt t l> X ' K X x y y x x X X WOOOOOt 10 » 10 it to tt «o «t to tt to GROWTH NCREMENTS f ROM PITH i n it >o n io tt to <t to tt to GROWTH INCREMENTS f ROM PITH K> It 10 » 10 It >0 4t 10 It 10 GROWTH INCREMENTS f ROM PITH Figure 4a. Scatter plot of height over nunber of growth increments at which juvenile - nature wood transition occurs. Figure 4b. Scatter plot of height over nunber of growth increnents at the base of dead branches for all trees. Figure 4c. Scatter plot of height over nunber of growth increnents which indicates the lowest relative density value in the profile. 47.5-45-42.5-40-37.5 35 32.5-30-27.5-X 25-o UJ 22.5-I 20-17.5-15-12.5-10 -7.5-5-2.5-0 y • -0.1313 «• 0.5833 X r • 0.8894 .8' 9' • D • 12.5 15 17.5 20 22.5 75 27.5 30 32.5 35 37.5 40 42.5 45 47.5 TOTAL HfTGHT (m) Figure 3*. Scatter plot and height prediction aodel for crovn base height over total height. Figure 3b. Scatter plot and height prediction aodel for average crovn height over total height. 116 47.5-: i i 1 1 1 1 1 1 1 ! : 1 1 1— 12.5 15 17.5 20 22.5 25 27.5 30 32.5 35 37.5 40 42.5 45 47.5 TOTAL HEIGHT (m) Figure 6. Scatter plot of a) total height and height prediction todels over total height; for: b) average crown height; and c) crown base height. 117 4i s-i i i ' i , 1 i i 1 i i \ .* , 0 5 » 15 30 71 JO ii 40 45 50 55 GROWTH INCREMENTS FROM PITH Figure 7. Graphical representation of: a) total tree height; and b) crovn base height over nunber of growth increnents fron the pith at breast height. 118 4J.S-GROWTH INCREMENTS FROM PITH Figure 8. Graphical representation of: a) total tree height; b) crovn base height before harvest; c) live crovn base height at harvest; and d) average crovn height at harvest over nuaber of growth increaents froa the pith at breast height. Figure Sa. Scatter and height prediction nodel for total tree height over nunber of growth increments froi the pith at breast height. V = -3.5855 + 0.9854x - 0.0037x r = 0.9324 x x X X X . xy/y. ,'X x X /x »- X XX? XXX 10 It 70 25 SO SB 40 45 50 B5 60 GROWTH INCREMENTS FROM PITH Figure 9b. Scatter and height prediction nodel for lowest relative density over nuiber of growth increnents fron the pith at breast height. 2 y = -6.3482 + 1.0073x - O.OO88X r = 0.9089 40 6 39-37.5-38 34.5-33-31.6-30-28 6-27-? 26.5 J '•I'l 1 i I 1 i i i i 0 6 10 16 20 25 30 35 40 46 60 65 60 GROWTH INCREMENTS FROM PITH 46-43 5 42 Figure 10a. Scatter and height prediction nodel for height to crown base over nunber of growth increments froi the pith at breast height. 46 43 6 42 40 6 39 376 36 34 6 33 316 30 26 6 27 "E 266 t— 24 5 22 8 UJ X 2' 166 18 16 5 16 13 6 12 10 5 H 9 7.6 6 4.6 6.9988 + 0.4853X • 0.7 174 B B B BH B B p / / / / / / / / / / B J» »»*B BEE B BB 10 16 20 26 SO 36 40 46 60 65 80 GROWTH INCREMENTS FROM PITH Figure 10b. Scatter and height prediction nodel for juvenile - nature wood transition points over nunber of growth increments fron the pith at breast height. 121 1 i 11 -i—1—i 1 1 •! -i 'l 11 r——r-0 5 10 15 20 25 30 35 40 45 60 65 60 GROWTH INCREMENTS FROM PITH Figure 11. Height prediction nodels for: a) total tree height; b) lowest relative density height; c) crovn base height; d) juvenile - nature wood transition height over nunber of growth increnents fron the pith at breast height; and e) diagranatic tree representations. u z I o ie- y = 11.0912 - 0 4264x o r = -0.5646 16- o 14 - o 0 12- o o_ 10-o'~'-~- o o o o 8 o o *• * « ~ « ^ O 6- § ° 0 O 4 -o 2-Q n -2--4- O C 8 10 12 14 TRANSITION HEIGHT H Figure 12a. Scatter and height prediction aodel for the difference between crovn base height and juvenile - nature vood transition height over juvenile - nature vood transition height. 28-26 24-22-20 18-16-14 -12-10-8-6-4 -2-o o o 8 o 8 o y = 19.3270 - 0.2319x r = -0.2319 8 o o o <9 S 8 10 12 14 TRANSITION HEIGHT M —t— 16 Figure 12b. Scatter and height prediction aodel for the difference betveen total height and juvenile - nature vood transition height over juvenile - nature vood transition height. 123 Figure 13. Graphical representation of: a) estimation of the average transition height A; and b) estination of the average transition height height 8 over nunber of growth increnents froa pith at breast height. 124 41 46-44 42-40-16-16 14 c E S »H so .E .e *» E " U « < "* 14 12 H 10 • e 4-I 0 V\ *- >A — • r«> \ / \ / o —r— B.H —r~ 20%. T Legend A TREC-U5 X mt-u? • TR£[_-166 E TREE —1E11 E TREE-1C1 ¥ 1REJ^IC6 4- IfCE-IM & 1RCE-1D6 o Ifrr^iES, 4 TREC-1C6 O T«£-ir7 E TREE-iriO V 1REE-CR1 E 1BIC-CR? 40% AVG TRANS ( A ) TREE SECTION Figure 14. Scatter plot of juvenile - nature (J. N.) wood transition nuiber of growth increments over tree sections and overall section averages, i.e. average transition age A. 125 z UJ g LU X z o IS) Z < o 46 43 6-42-40 6-3* 37.5 se 34 5 33 315 30 235 27 26 6 24 22.6 21 18 6-18-16 6-15 13 6-12-106 • 76 e 4 6 * o •-r, Leoend TRCt -1*5 X met -U7 D 1BCC -»6 E iRtr -ten E mt -IC 1R£E -1C6 * If" e -106 0 IRCt •t -ue o mr -V7 c 1REE -irio V 1«£ -C»l E l«t -CK2 —r-B.H. 20% TREE SECTION —I 1 40% AVG TRANS (B) Figure IS. Scatter plot of height differences atong total and transition heights over tree sections and overall section averages, i.e. average transition height B. 47.5 45-42.3-40-37.3-35 32 5-30-(m) 27.5-y-X 25-O U 22.5-I 20 17 5-15 12.5-10-7.5-5-2.S-0-y = -5.1764 + 0.7894 X r = 0.8248 B B / a / / ^ B3 s B 1 I I I I 1 I I I I I ! I I I I2.S IS 17.5 20 22.5 23 27.5 30 32 3 33 37.5 40 42 3 43 47.5 TOTAL HEIGHT (m) Figure 16a. Scatter plot and height prediction nodel for average transition height A over total height. 47.5-45-42 5 40-37.3 33-32.3-30-(m) 27.5-23-X o UJ 22 5-X 20-17.5-15 12.5-10-7.5-5-2.5-0 \-y = -5.2892 + 0.6421 X r = 0.7811 ro x*.>* 17 5 15 17 5 20 22.5 25 27.5 30 32 5 33 37.3 40 42.3 43 47 5 TOTAL HEIGHT (m) Figure 16b. Scatter plot and height prediction nodel for average transition height B over total height. 127 Figure 17. Scatter plot of a) total tree height, and height prediction aodels over total height for: b) average crovn height; c) average transition height A; d) live crovn base height; and e) average transition height B. 128 GROWTH INCREMENTS FROM PITH Figure 19. Pith to hart relative density profiles for pruned tree RF3. 129 Figure 18. Pith to bark relative density profiles for pruned tree Rf2. 130 Appendix 1. Average Crown Height Estimation Procedure 1. The length of each branch (BL) was plotted against the vertical distance from the terminal leader to the base of the branch (L) to observe the actual shape of each sampled crown (Appendix la). 2. The length of each branch (BL) was plotted against the transformed vertical distance from the terminal leader to the base of the branch (Appendix lb): ln [ ( L/c ) + 1 ] where: ln = natural logarithm L = distance from the terminal leader to the branch base or crown 1ength c = coefficient that describes the curvature of the crown profile (estimated for Douglas-fir as 6.1 by Mitchell (1975)) 131 The slope of each crown, or regression coefficient (b) values that relate branch growth to height growth were estimated using the following regression equat i on: BL = bd [ In ( L/c ) + 1 ] (1) where: BL = branch length b = regression coefficient that relates branch growth to height growth d = coefficient that compensates for branch crooks (estimated for Douglas-fir as 0.975, by Mitchel1 (1975)) The average crown radius, which represented the maximum branch length (BL), was utilized to estimate the distance from the terminal leader to the crown base using Equation 1 and solving for L (Appendix lc) as fol1ows: L = c ( e BL/(bd) - 1 ) where: e = 2.71828 Average crown height was calculated by subtracting L from the total tree height (Table 3). 132 • i i i i ' i i i I 0 1.5 3 4.5 6 7.5 9 10.5 12 13.5 VERTICAL DISTANCE FROM THE LEADER (L) (m) Appendix la. Relationship between total branch length (BL) and the vertical distance froi the leader (L) for Satple Tree 1A5. 133 4.5 3.5-X o z X u z < cr m 2.5-0.5 0 0.2 0.4 0.6 0.8 1 1.2 1.4 1.6 VERTICAL DISTANCE FROM THE LEADER ln[(L/c)+l)] Appendix lb. Relationship between total branch length (BL) and the transforaed vertical distance froi the leader ln [ (L/cHl ] for Saiple Tree 1A5 . 134 - A: CROWN RADIUS J Q' o/ O / 4 7 - 0 o/ 0 0 w 0 / 0 4 -0 4 -0 4 /o / 0 c 0 0 0 .6 0 0 0 0 4 0 t 0 0 U-—i— -i 1 1 -4 B: AVERAGE CROWN POSITION 0-2 0.4 0.6 0.6 Bl 1.2 1.4 1.6 VERTICAL DISTANCE FROM THE LEADER ln[(L/c)+l)] Appendix lc. Average crown height position determination for saaple tree 1A5. 135 BRANCH YEARS Appendix 2. Relationship between total height and branch age to estimate base of live crown positions at young ages. 136 Appendix 3. Summary of X-Ray Densitometric Analysis Procedure 1. After air drying, the wood sample strips were mounted and glued between two mounting sticks. 2. The mounted samples were reduced to a uniform thickness of two mm using a specially designed twin blade circular saw. 3. Identification characters were then written on the mounts adjacent to the wood samples using an x-ray opaque lead based paint. 4. The wood samples were extracted by immersion in a solution of 1:2 alcohol benzene for 30 minutes and then air dried at room temperature. 5. A portion of the core mounting stick was removed to measure the angle between the longitudinal tracheids and the long axis of the sample surface. This angle was measured with a goniometer in the eyepiece of a low power stereoscopic microscope. 6. The wood samples along with calibration wedges were placed on fine grained, high resolution, single emulsion x-ray films. The calibration wedges, made of Douglas-fir wood blocks of known relative density, were utilized in the conversion of film density into wood relative density data. 137 7. Radiographs were then made by exposing the wood samples, calibration wedges and x-ray films on a moving col 1imated x-ray scanning machine oriented according to the angle of the longitudinal tracheids. 8. The radiographs were developed in specially designed film processing tanks containing separate solutions of developer, stop bath indicator and fixer with hardener. The tanks were placed in a temperature controlled waterjacket. The length of time and temperature at which the radiographs were exposed to the chemicals was determined according to the chemical manufacturer's instructions. 9. The developed radiographs were examined on a light table under a low power stereoscopic microscope. The pith date and the corresponding growth increment calendar years were marked, in decades, on the films. 10. The radiographs were placed on a computerized scanning densitometer that converted the wood sample image on the film into growth increment width and relative density data, measured at 0.01 mm and 0.05 mm steps along the radius of the wood sample, respectively. 11. The data obtained from the densitometer were stored on magnetic tapes for further processing and summarizing. Forintek Canada Corporation's data acquisition Tree Ring Input Program (TRIP) was used to obtain the average relative density of each growth i ncrement. 138 Appendix 4. Fortran prograi to deternine residual sua of •quarts using non-linear optimization routines for segnented regression models. List mg of FAST 1 $R •FTN SCARDS'DAVID S SPR1NT--C 2 R -LOAD*NA:NLMON 3 ASSIGN 1-1A53RD<8) 4 CALL GETDAT 5 IN P 6 .41025..35647E-2.20. .4924 1E-2 7 PR F B EX SIMPLX 9 1 10 200.10 11 IE-05 12 EX FNMIN 13 200 14 10.1.E-11 16 STOP 17 SET COST-ON Listing of ...•DAVID.S 1 FUNCTION XDFUNC(X.N) 2 IMPLICIT REAL'S!A-H.O-Z) 3 COMMON /SHARE/POINTS(2. 100).NDAT 4 REAL'S XfN) 5 XDFUNC'0.0 6 DO 100 I-1.NDAT 7 IF(POINTS(1.1) GT X(3) )XOFUNC«XDFUNC+(POINTS!2.I)-B 1 (X!1)+X(2)'X(3)*X(4)'(P0INTS(1.I)-X!3))))"2 9 IF(POINTS(1,l).LE.X(3))XDFUNC"XDFUNC*(POINTS!2, I ) -10 1 <X(1)*X(2)'P0INTS(1.I)))"2 11 100 CONTINUE 12 RETURN 13 END 14 C 15 C 16 C 17 SUBROUTINE GETDAT 18 IMPLICIT REAL'S!A-H.O-Z) 19 COMMON /SHARE/P0INTS(2.100).NDAT 20 DO 100 1-1.100 21 READ(1.2OO.END»5OO)P0INTS(1.I).POINTS!2.I) 22 20O FORMAT(T6.F2.0.F11.7) 23 WRITE(6.2OO)P0INTS(1.I).POINTS(2.I) 24 NDAT-I 25 100 CONTINUE 26 500 RETURN 27 END 1 |HEIGHT-803? HEIGHT-60% 0 6-06-0 4 - lHEIGHT-40%1 06 0 6-0.4 1 I 1 1 i 1 I 1 I 1 I -5 10 15 20 25 30 35 40 45 50 55 GROWTH INCREMENTS FROM PITH Appendix S. a) Tree IAS. Simple and segaented linear regression models on pith to bark relative density profiles. Legend A HT/C. INCREMENTS TROM PITH X HT/LOWEST RELATIVE DENSITY • HT/CROWN BASE EJ HT/J.- M. WOOD TRANSITION _ I i I 1 I I 1 1 I 1 II.' 0 6 10 16 20 25 30 35 40 46 60 66 60 GROWTH INCREMENTS FROM PITH Appendix 5. b) Tree IAS. Relationships between total tree height, lowest relative density height, crown base height and juvenile - mature wood transition over nuaber of growth increments from pith at breast height. 1 1 1 1 1 1 i 1 1 1 r 5 10 15 20 25 30 35 40 45 50 55 GROWTH INCREMENTS FROM PITH Appendix f>. a) Tree 1A7. Siapie and segaented linear regression aodels on pith to bark relative density profiles. Legend A HT/G. INCREMENTS FROM PITH X HT/LOWEST RELATIVE DENSITY • HT/CROWN BASE H HT/J - M. WOOD TRANSITION -1 1 I 1 1 I 1 1 1 1 I 1—1 0 5 10 15 20 25 30 35 40 45 60 65 60 GROWTH INCREMENTS FROM PITH Appendix f>. b) Tree 1A7. Relationships between total tree height, lowest relative density height, crown base height and juvenile - nature wood transition over nuaber of growth increaents froa pith at breast height. 10 15 ?0 75 Jf 35 40 «5 50 Si GROWTH INCREMENTS FROM PITH Appendix 7. a) Tree IBS. Sinple and segnented linear regression nodels on pith to bark relative density profiles. i i {iii 1 1 i 1 1 1 i i 0 6 10 16 20 25 30 36 40 46 60 65 60 GROWTH INCREMENTS FROM PITH Appendix 7. b) Tree 1B6. Relationships between total tree height, lowest relative density height, crown base height and juvenile - nature wood transition over nunber of growth increnents fron pith at breast height. 0.6 0 5^ 0.4 0.6-0.5 0 4 tn 0.6 z 0.5 0 4-0 6 06 0 4 0.6 0 6-1 0.4 lHEIGMT-,B0%j [HEJGHT-60*J |HEIGHT-20% HEIGHT-B.H. 5 10 15 70 ?5 ?0 J5 40 45 50 55 60 GROWTH INCREMENTS FROM PITH Appendix 8. a) Tree 1BU. Siaple and segaented linear regression aodels on pith to bark relative density profiles. 46 435 42 40.6 39 37.6 36-34 6 33 31.5 30-28 6 27-25 6-24 Legend A HT/G. INCREMENTS FROM PITH X HT/LOWEST RELATIVE DENSITY • HT/CROWN BASE  HT/J.- M. WOOD TRANSITION_ •P-N3 15 20 25 30 35 40 45 50 GROWTH INCREMENTS FROM PITH Appendix 8. b) Tree IBM. Relationships between total tree height, lowest relative density height, crown base height and juvenile - nature wood transition over nunber of growth increments froa pith at breast h*ight. 0.6-0 5-04 0 6 0 5 04 06 06 1 04 06 06 0-4 H 0 6-t 06 04 [HEIGHT-8 0*| [HEIGHT-60%] |HEIGHT-40%J |HEIGHT-20lt] |HEIGHT-B.H"] —i . i IIIIIIII— 5 10 15 20 25 JO 35 40 45 50 55 GROWTH INCREMENTS FROM PITH Appendix 9. a) Tree 1C1. Sinple and segnented linear regression nodels on pith to bark relative density profiles. 46-43 6-42-40.6-39-37.6-36-34 6 -33-316-30-28 6-27-(m) 26.5-r— 24-X 22 5-o UJ 21-19.5-18-16.6-15-13.6-12-10.6 9 -7.6-6-4.6-3-1.6 A Legend A HT/G. INCREMENTS TRQM PITH X HT/LOWEST RELATIVE DENSITY • HT/CROWN BASE HT/J.- M. WOOD TRANSITION —T— 10 T I 1 1 1 1— 20 25 30 35 40 46 50 GROWTH INCREMENTS FROM PITH -1— 15 55 60 Appendix 9. b) Tree 1C1. Relationships between total tree height, lowest relative density height, crown base height and juvenile - nature wood transition over nunber of growth increnents fron pith at breast height. I I i 1 I r 10 15 70 75 ™ J5 40 45 50 55 GROWTH INCREMENTS FROM PITH Appendix 10. a) Tree 1C6. Siaple and segaented linear regression models on pith to bark relative density profiles. 45-43 5 42 40.5 30 37.5-36 34.6-33-31.6-30-28 5-27-25.6 24 22 5 2H 19.6 18 16.6-15 13 6-12 10.5 9 7.5 6 4.6 3 1-8 A Legend A HT/G. INCREMENTS FROM PITH X HT/LOWEST RELATIVE DENSITY • HT/CROWN BASE 0 y_TZLli*- WOOD TRANSITION -r i i 1 i 1 i 1 i I 10 15 20 25 30 35 40 45 60 55 60 GROWTH INCREMENTS FROM PITH Appendix 10. b) Tree ICS. Relationships between total tree height, lovest relative density height, crovn base height and juvenile - nature vood transition over nuaber of growth increments froa pith at breast height. 0.6 OS 0 4 0.4- ••• ••• [HEIGHJ-80») IHEIGHT-B.'HTI 5 10 15 2n 75 30 35 40 45 50 55 GROWTH INCREMENTS FROM PITH Appendix 11. a) Tree IDS. Sinple and segnented linear regression nodels on pith to bark relative density profiles. Legend A HT/G. INCREMENTS FROM PITH X HT/L0WE5T RELATIVE OENSITY • HT/CROWN BASE  HT/J.- M. WOOD TRANSITION 1— 15 20 25 30 35 40 GROWTH INCRFMENTS FROM PITH 46 -I— 60 -r— 65 —r-60 Appendix II. b) Tree IDS. Relationships between total tree height, lowest relative density height, crown base height and juvenile - nature wood transition over nunber of growth increnents fron pith at breast height. HEIGHT-80%, [HEJGHT-60%; HEIGHT-40% HEIGHT-20% |HEIGHT-BR1 1 1 i 1 : 1 1—1 1 1— 5 10 15 70 75 JO J5 <n A* 50 55 GROWTH INCREMENTS FROM PITH Appendix 12. a) Tree 1D6. Staple and segaented linear regression aodels on pith to bark relativt density profiles. Legend A HT/G. INCREMENTS FROM PITH X HI/LOWEST RELATIVE DENSITY • HT/CROWN BASE B HT/J.- M. WOOD TRANSITION / »A X J , 'SI -1— 15 70 25 30 35 40 45 60 GROWTH INCREMENTS FROM PITH Appendix 12. b) Tree ID6. Relationships between total tree height, lowest relative density height, crown base height and juvenile - aature wood transition over nunber of growth increments froa pith at breast height. —i— 40 10 65 60 06 05 0 4 0.6 0.6 0 4 >-*-in 0 6 z UJ ui 0.6 5 04 ui EE 06 06 0 4 0.6-1 0 6 0.4-V HEIGHT-80% |HEtGHT-40tt| |HEIGHT-20%| |HEIGHT-B"H"1 4n sn 55 GROWTH INCREMENTS FROM PITH Appendix 13. a) Tree 1E3. Sinple and segnented linear regression nodels on pith to bark relative density profiles. 15 20 25 30 35 40 45 60 GROWTH INCRFMENTS FROM PITH Appendix 13. b) Tree 1E3. Relationships between total tree height, lowest relative density height, crown base height and juvenile - nature wood transition over nunber of growth increnents fron pith at breast height. i i 1 '• i I 1 I 1 1 5 10 15 70 25 30 35 40 45 50 55 GROWTH INCREMENTS FROM PITH Appendix 14. a) Tree 1E8. Simple and segaented linear regression aodels on pith to bark relative density profiles. Legend A HT/C. INCREMENTS FROM PITH X HT/LQWEST RELATIVE DENSITY • HT/CROWN BASE B HT/J.- M. WOOD TRANSITION GROWTH INCREMENTS FROM PITH lix 14. b) Tree IE8. Relationships between total tree height, lowest relative density height, crovn base height and juvenile - aature vood transition over nunber of growth increments froa pith at breast height. 0 6 -05 0.4 -0.6-05 0 4 V) 0.6 ui OS l 0.4 UJ K 06 0.6 0.4 06 06 0.4 A-[HEIGHT-80%! I^EIGHT-60%J I HEIGHT-40%) HEIGHT-20% |HEIGHT-BH1 5 10 15 70 75 JO 35 40 45 50 55 GROWTH INCREMENTS FROM PITH Appendix IS. a) Tree 1F7. Sinple and segnented linear regression nodels on pith to bark relative density profiles. 1 I I 1 I 10 15 20 26 30 35 40 45 50 55 60 GROWTH INCREMENTS FROM PITH Appendix 15. b) Tree 1F7. Relationships between total tree height, lowest relative density height, crown base height and juvenile - nature wood transition over nunber of growth increnents fron pith at breast height. 06 0.5-0 4-0 6-0 5 0 4 ui 0.6 z 0.5 1 0.4 06 0.6 0.4 0.6-06 0.4 H [HEIGHT-80%J |HEIGHT-40»1 |HEIGHT-20%| |HEIGHT-B.H~1 5 10 t!> 70 75 30 35 40 45 50 55 GROWTH INCREMENTS FROM PITH Appendix 16. a) Tree 1F10. Siaple and segaented linear regression aodels on pith to bark relative density profiles. 45-43.6-42-40.6-38-37.6-I I I i I i 1 1 1 ill1 0 6 10 15 20 25 30 35 40 45 SO 65 60 GROWTH INCREMENTS FROM PITH Appendix 16. b) Tree IF10. Relationships between total tree height, lowest relative density height, crown base height and juvenile - nature wood transition over nunber of growth increments froa pith at breast height. Legend A HT/G. INCREMENTS FROM PITH X HT/LOWEST RELATIVE DENSITY • HT/CROWN BASE G3 HT/J.- M. WOOD TRANSITION --[HEIGHT-80%| <fv. |HEIGHT-40%] 1 HEIGHT-20% 1 1 1 i 1 1 i 1 [HEIGHT-B.HJ GROWTH INCREMENTS FROM PITH Appendix 17. a) Tret CR1. Sinple and segnented linear regression nodels on pith to bark relative density profiles. Legend A HT/G, INCREMENTS FROM PITH X HT/LOWEST RELATIVE DENSITY B HT/J.- M. WOOD TRANSITION I i i 1 i 1 : 1 1 i 1 1 1 • 0 6 10 15 20 25 30 35 40 45 60 65 60 GROWTH INCREMENTS FROM PITH Appendix 17. b) Tree CRI. Relationships between total tree height, lowest relative density height, crown base height and juvenile - nature wood transition over nunber of growth increnents fron pith at breast height. 5 m 15 20 25 JO J5 40 4b 50 55 GROWTH INCREMENTS FROM PITH Appendix 18. a) Tree CR2. Sinple and segmented linear regression aodels on pith to bark relative density profiles. Legend A HT/G. INCREMENTS FROM PITH X HT/LOWEST RELATIVE DENSITY H HT/J.- M. WOOD TRANSITION 1 I 1 I I I 1 1 1 1 1 1 0 6 10 15 20 25 30 35 40 45 50 65 60 GROWTH INCREMENTS FROM PITH Appendix 18. b) Tree CR2. Relationships between total tree height, lowest relative density height, crown base height and juvenile - nature wood transition over nuaber of growth increnents froa pith at breast height. 0.6 |HEIGHT-80%| |HEIGHT-60%| HEIGHT-40%| \A :.... HEIGHT-20%| • • ! • HEIGHT-B H.| 1 1 I 1 1 1 1 1 I 1 r-5 10 15 20 25 SO 35 40 45 50 55 GROWTH INCREMENTS FROM PITH Appendix 19. a) Tree RFt. Sinple and segaented linear regression aodels on pith to bark relative density profiles. 46 43.5 42 40.6 39 37.6 36 34.6 33 31.5 30 286 27 25.5 24-22.6 21 19.6 18-16 5 16 13 6 12-10.6 9-7.6 8 4.6 3 1-6 A Legend A HT/G. INCREMENTS FROM PITH X HT/LOWEST RELATIVE DENSITY • HT/CROWN BASE HEIGHT 10 16 20 25 30 36 40 45 50 GROWTH INCREMENTS FROM PITH —i i— 65 60 Appendix 19. b) Tree RF1. Relationships between total tree height, lowest relative density height and crown base height over mincer of growth increments froa pith at breast height. 154 Appendix 20. Sunnary of relative density data. a) Tree IAS Height Oianeter Section Nunber of 6rovth Relative Density (n) (cn) Height Increnents I.B. O.B. Juv. Hat. Total Juvenile Hood Nature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 46.8 40.3 B.H. 22 34 56 .486 .058 .620 .380 .572 .044 .640 .480 .542 .062 .640 .380 7.5S 39.3 33.8 201 28 19 47 .438 .054 .570 .360 .513 .044 .620 .420 .468 .063 .620 .360 14.90 34.1 29.5 40X 25 15 40 .422 .043 .540 .370 .483 .033 .520 .400 .445 .049 .540 .370 22.20 27.0 25.9 601 28 - 28 .460 .038 .560 .400 - - - - .460 .038 .560 .400 29.40 19.5 18.7 BOX 13 - 13 .440 .030 .510 .410 - - - - .440 .030 .510 .410 b) Tree 1A7 Height Dianeter Section Nunber of firovth Relative Density (n) (cn) Height Increnents I.B. O.B. Juv. Nat. Total Juvenile Hood Nature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 46.1 39.8 B.H. 19 32 51 .525 .052 .620 .430 .597 .040 .660 .460 .569 .057 .660 .430 7.20 41.7 37.3 201 24 20 44 .482 .067 .600 .370 .567 .042 .650 .470 .520 .071 .650 .370 15.10 37.0 32.8 401 24 IS 39 .471 .046 .570 .400 .495 .042 .550 .380 .480 .045 .570 .380 22.50 30.7 26.5 601 31 - 31 .479 . 054 . 670 . 410 - - - - .479 . 054 . 670 . 410 29.90 17.6 16.1 801 18 - IB .499 . 038 . 600 . 450 - - - - .499 . 038 . 600 . 450 155 Appendix 20. Suaaary of relative density data (cont.l. c) Tree 116 Height Diameter Section Number of Growth Relative Density (m) (cm) Height Increments l.B. O.B. Juv. Hat. Total Juvenile Hood nature Mood Total Hood Wood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 49.2 40.0 B.H. 16 37 S3 .467 .035 .560 .420 .551 .042 .650 .450 .520 .055 .650 .420 8.SS 39.S 36.S 201 17 28 45 .448 .038 .S60 .400 .539 .030 .590 .490 .504 .OSS .590 .400 17.20 33.5 31.5 401 18 14 32 .470 .034 .540 .420 .526 .027 .570 .490 .495 .042 .570 .420 25.80 25.6 24.5 601 24 - 24 .478 .027 .540 .440 - - - - .478 .027 .540 .440 36.10 14.5 13.3 802 12 - 12 .479 .052 .610 .420 - - - - .479 .052 .610 .420 d) Tree 1B11 Height Diameter Section Number of Growth Relative Density (m) (ca) Height Increments l.B. O.B. Juv. Hat. Total Juvenile Hood Nature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 49.1 43.5 B.H. 32 25 63 .515 .044 .610 .430 .572 .041 .680 .510 .543 .057 .680 .430 7.10 41.8 37.7 201 25 28 53 . 480 . 039 . 570 . 410 . 4% .034 .590 .440 .489 .037 .590 .410 14.20 36.5 32.0 401 16 29 45 .479 .032 .530 .410 .477 .034 .580 .430 .478 .033 .580 .410 21.30 27.7 24.7 601 29 - 29 .479 .031 .560 .430 - - - - .479 .031 .560 .430 28.45 15.8 14.0 80Z 19 - 19 .488 .032 .570 .450 - - - - .488 .032 .570 .450 156 Appendix 20. Sunnary of relative density data (cont.). e) Tree IC1 Height Dianeter Section Nunber of Growth Relative tensity (n) (cn) Height Increnents I.B. O.B. Juv. Nat. Total Juvenile Hood Hature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 47.3 40.7 B.H. 19 35 54 .503 .037 .590 .440 .586 .035 .660 .490 .558 .053 .660 .440 8.00 40.1 36.2 201 25 22 47 . 464 .043 . 560 . 390 . 530 . 027 . 570 . 470 . 490 . 049 . 570 . 390 16.00 36.1 32.9 401 29 10 39 .458 .030 .520 .400 .497 .019 .520 .450 .468 .032 .520 .400 24.15 29.8 25.9 601 29 - 29 .448 .023 .510 .410 - - - - .448 .023 .510 .410 32.00 12.0 14.9 80Z 15 - 15 .411 .015 .460 .400 .... .411 .015 .460 .400 f) Tree 1C6 Height Dianeter Section Nunber of firovth Relative tensity (n) (ci) Height Increnents I.B. O.B. Juv. Hat. Total Juvenile Hood Hature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 47.1 37.5 B.H. 15 38 S3 .502 .052 .650 .440 .592 .046 .700 .500 .566 .062 .700 .440 7.60 39.8 34.2 201 35 11 46 .489 .059 .580 .380 .544 .032 .600 .510 .502 .059 .600 .380 15.20 35.1 31.5 402 26 10 36 .488 .043 .560 .430 .518 .019 .560 .500 .496 .040 .560 .430 22.80 29.0 26.7 601 25 - 25 .478 .034 .550 .410 - - - - .478 .034 .550 .410 30.40 14.4 13.2 801 12 - 12 .499 .035 .560 .430 - - - - .499 .035 .560 .430 157 Appendix 20. Sunnary of relative density data (cont.). g) Tree 10S Height <n) Dianeter (cn) I.B. O.B. Section Height Nunber of 6routh Increnents Juv. Hat. Total Hood Hood Sect. Relative Density Juvenile Hood Hean S.D. Hax. Hin. Hature Hood Hean S.D. Hax. Hin. Total Hean S.D. Hax. Hin. 1.30 45.5 26.8 B.H. 13 14 33 .462 .031 .540 .420 .490 .032 .540 .440 .474 .034 .540 .420 6.40 35.7 31.9 201 20 7 27 .466 .026 .500 .430 .530 .032 .560 .480 .482 .039 .560 .430 12.80 27.6 25.9 401 20 - 20 .475 .035 .550 .430 - .475 .035 .550 .430 19.20 20.3 18.2 60Z 14 - 14 .456 .034 .510 .390 - .456 .034 .510 .390 25.60 11.3 10.1 801 8 - 8 .540 .027 .580 .500 - .540 .027 .580 .500 h) Tree 1D6 Height Dianeter Section Nunber of firovth Relative Density (n) (cn) I.B. O.B. Height Increnents Juv. Hat. Total Hood Hood Sect. Juvenile Hood Hean S.D. Hax. Hin. Nature Hood Hean S.D. Hax. Hin. Total Hean S.D. Hax. Hin. 1.30 43.2 38.5 B.H. 17 18 35 .445 .055 .550 .380 .530 .033 .600 .490 .488 .062 .600 .380 5.90 .384 34.3 201 15 14 29 .407 .031 .470 .350 .468 .031 .520 .430 .437 .043 .520 .350 11.90 33.4 32.1 40Z 23 - 23 .450 .037 .550 .400 - .450 .037 .550 .400 17.70 25.8 23.4 601 17 - 17 .428 .041 .530 .360 - .428 .041 .530 .360 23.60 13.2 12.2 801 8 8 .468 .027 .510 .420 _ _ - _ .468 .027 .510 .420 158 Appendix 20. Summary of relative density data (cont.). i) Tree 1E3 Height Diameter Section Number of 6rovth Relative Density (m) (cm) Height Increments l.B. O.B. Juv. Hat. Total Juvenile Hood Nature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Mean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 39.4 37.6 B.H. 28 15 43 .574 .045 .670 .500 .634 .028 .680 .590 .595 .049 .680 .500 7.10 35.3 32.5 201 26 6 32 .487 .030 .560 .440 .546 .031 .590 .500 .498 .038 .590 .440 14.00 31.3 28.5 401 11 15 26 .444 .034 .510 .390 .454 .013 .480 .430 .450 .024 .510 .390 21.00 22.8 21.1 601 17 - 17 .417 .017 .450 .400 - - - - .417 .017 .450 .400 28.00 12.8 11.4 80Z 9 9 .483 .031 .560 .450 - - - - .483 .031 .560 .450 j) Tree 1E8 Height Diaaeter Section Number of 6rovth Relative Density (a) (ca) Height Increments l.B. O.B. Juv. Hat. Total Juvenile Hood Nature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 68.2 57.1 B.H. 34 13 47 .472 .042 .590 .390 .527 .016 .570 .510 .487 .044 .590 .390 7.20 54.7 48.6 201 26 IS 41 .441 .057 .610 .360 .503 .026 .570 .460 .463 .057 .610 .360 14.40 45.4 40.5 40Z 33 - 33 .456 .039 .530 .370 - - - - .456 .039 .530 .370 21.60 36.3 32.5 601 25 - 25 .420 .034 .500 .360 - - - - .420 .034 .500 .360 28.80 21.3 19.7 601 14 - 14 .454 .030 .530 .420 - - - - .454 .030 .530 .420 159 Appendix 20. Summary of relative density data (cont.). k) Tree 1F7 Height Dianeter Section Nunber of Growth Relative Density (•) (cn) Height Increments l.B. O.B. Juv. Hat. Total Juvenile Hood Nature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 S7.S 48.3 B.H. 25 28 53 .503 .053 .650 .430 .595 .025 .640 .550 .551 .061 .650 .430 8.30 47.3 44.0 20Z 25 21 46 .454 .061 .610 .380 .533 .026 .600 .510 .490 .061 .610 .380 16.40 39,9 37.3 401 24 14 38 .470 .043 .570 .420 .478 .033 .550 .430 .473 .040 .570 .420 24.90 30.9 26.8 601 22 - 22 .497 .056 .650 .430 - - - - .497 .056 .650 .430 33.70 16.1 14.7 801 14 - 14 .517 .058 .640 .450 - - - - .517 .058 .640 .450 1) Tree 1F10 Height Diaaeter Section Nuaber of Growth Relative Density (a) (ca) Height Increaents l.B. O.B. Juv. Nat. Total Juvenile Hood Nature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 52.7 45.3 B.H. 24 28 52 .492 .054 .640 .430 .576 .030 .630 .510 .537 .060 .640 .430 7.80 41.S 36.4 207. 22 24 46 .450 . 048 . 570 . 390 . 517 . 032 . 570 . 450 . 485 . 052 . 570 . 390 15.90 35.0 31.3 401 15 24 39 .446 .028 .490 .410 .487 .021 .530 .440 .472 .030 .530 .410 23.40 28.0 23.9 601 26 - 26 .495 .026 .550 .460 - - - - .495 .026 .550 .460 31.20 16.5 14.4 801 19 - 19 .470 .042 .560 .430 - - - - .420 .042 .560 .430 160 Appendix 20. Sunnary of relative density data (cont.). n) Tree Cfil Height Dianeter Section Nunber of Growth Relative Density (n) (cn) Height Increnents ~ I.B. O.B. Juv. Hat. Total Juvenile Hood Hature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Max. Hin. Hean S.D. Hax. Hin. 1.30 22.1 20.0 B.H. 20 17 37 .491 .054 .610 .410 .590 .046 .670 .520 .536 .070 .670 .410 3.34 19.8 18.4 201 19 15 34 .490 .054 .660 .420 .612 .034 .680 .560 .543 .076 .680 .420 5.10 18.0 16.8 40Z 31 14 31 .486 .069 .620 .390 - .486 .069 .620 .390 9.95 12.3 11.4 601 17 - 17 .485 .025 .540 .450 - .485 .025 .540 .450 13.36 6.3 5.7 801 7 - 7 .485 .040 .540 .430 - .485 .040 .540 .430 n) Tree CR2 Height Dianeter Section Nunber of 6rowth Relative Density (n) (cn) Height Increnents I.B. O.B. Juv. Hat. Total Juvenile Hood Nature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.40 42.4 38.4 B.H. 23 6.40 36.8 34.5 201 13 11.80 33.6 31.7 401 22 18.72 24.8 23.4 601 20 25.77 10.7 10.0 801 16 12 35 .428 .038 .510 .340 .473 16 29 .443 .037 .500 .370 .514 22 .488 .028 .540 .440 -20 .459 .048 .630 .400 -16 .493 .041 .590 .440 -.017 .500 .440 .444 .039 .510 .340 .032 .580 .440 .483 .049 .580 .370 - - - .488 .028 .540 .440 - .459 .048 .630 .400 - - - .493 .041 .590 .440 161 Appendix 20. Suanary of relative density data (cont.). o) Tree RF1 Height Dianeter Section Nunber of Growth Relative Density (a) (ca) Height Increaents l.B. O.B. Juv. Hat. Total Juvenile Hood Nature Hood Total Hood Hood Sect. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. Hean S.D. Hax. Hin. 1.30 28.3 26.0 B.H. 21 - 21 .429 .041 .530 .360 - - - - .429 .041 .530 .360 4.54 24.3 22.3 201 17 - 17 .423 .035 .500 .380 - - - - .423 .035 .500 .380 9.00 20.6 18.6 40Z 13 - 13 .398 .054 .500 .340 - - - - .398 .054 .500 .340 13.65 15.0 14.0 60Z 8 - 8 .445 .042 .510 .400 - - - - .445 .042 .510 .400 18.16 7.4 0.6 B0I 4 - 4 .492 .025 .520 .460 - - - - .492 .025 .520 .460 p) Tree RF2 Height (a) Dianeter (ca) l.B. O.B. Section Height Nuaber of Growth Increaents Juv. Hat. Total Hood Hood Sect. Relative Density Juvenile Hood Hean S.D. Hax. Hin. Nature Hood Hean S.D. Hax. Hin. Total Hean S.D. Hax. Hin. 1.30 59.4 68.3 B.H. 21 30 51 .425 .021 .480 .370 .475 .045 .640 .410 .454 .045 .640 .370 3.80 51.6 61.0 10Z 17 30 47 .427 .028 .490 .380 .489 .050 .620 .430 .466 .053 .620 .380 6.30 50.7 56.0 201 44 - 44 .421 .027 .490 .360 - .421 .027 .490 .360 12.30 44.4 50.3 40Z 37 - 37 .432 .026 .500 .390 - .432 .026 .500 .390 18.90 31.1 34.6 601 26 26 .425 .035 .560 .380 _ .425 .035 .560 .380 162 Appendix 20. Sunnary of relative density data (cont.). q) Tree Rf3 Height (n) Dianeter (cn) I.B. O.B. Section Height Hunber of firovth Increnents Juv. Hat. Total Hood Hood Sect. Relative Density Juvenile Hood Hean S.D. Hax. Hin. Hature Hood Hean S.D. Hax. Hin. Total Hean S.D. Hax. Hin. 1.30 37.5 44.0 B.H. 15 30 45 .482 .035 .600 .440 .523 .023 .580 .490 .509 .033 .600 .440 2.90 34.4 40.5 10Z 13 30 43 .438 .039 .560 .400 .490 .029 .570 .430 .474 .040 .570 .400 4.35 32.0 36.B 20Z 39 39 .458 .036 .580 .380 - .458 .036 .580 .380 11.60 25.0 30.4 401 33 33 .431 .026 .490 .380 - .431 .026 .490 .380 17.75 17.0 19.6 60Z 21 21 .470 .027 .560 .440 _ .470 .027 .560 .440 

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