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Dry-matter production and complete-tree utilization of lodgepole pine in Alberta Johnstone, Wayne David 1973

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15127 DRY-MATTER PRODUCTION AND COMPLETE-TREE UTILIZATION OF LODGEPOLE PINE IN ALBERTA by WAYNE DAVID JOHNSTONE B.S.F., U n i v e r s i t y of B r i t i s h Columbia, 1966 M.F., U n i v e r s i t y of B r i t i s h Columbia, 1967 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n the Department of FORESTRY We accept t h i s t hesis as. conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA A p r i l , 1973 I n p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f the r e q u i r e m e n t s f o r an advanced degree a t t h e U n i v e r s i t y o f B r i t i s h C olumbia, I agree t h a t the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r agree t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p urposes may be g r a n t e d by the Head o f my Department o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Department o f Forestry The U n i v e r s i t y o f B r i t i s h Columbia Vancouver 8, Canada Date April. 1973 i ABSTRACT Chairman: Dr. D. D. Munro Lodgepole pine (Pinus contorta Dougl. var. lat i f o l i a Engelm.) is a species of considerable importance to the forest economies of Alberta and the Interior of British Columbia. The objectives of the present thesis are: a. to present the results of studies of the dry-matter production, growth, and complete-tree utilization of 100-year-old lodgepole pine trees, b. to compare the yields of dry-matter from 100-year-old lodgepole pine stands grown under a variety of site and stand density conditions, and c. to compare the above-ground total standing crops of similarly aged lodgepole pine and Populus stands grown on similar sites. Tree and component weights of eighty-five, 100-year-old lodgepole pine trees from two average density stands and two-hundred and twenty-one, 100-year-old lodgepole pine trees from one high density stand located in southwestern Alberta were examined. Both graphical and regression techniques were used to develop allometric relationships between the component dry-weights and some easily measured tree characteristics. Of the independent variables tested, the combined variable of tree diameter at breast height squared times total tree height (D2H) was most closely associated with the component dry-weights. Reliable estimates of tree and component weights were obtained using the aforesaid independent variable. The systematic errors (under-estimates) resulting from the use of logarithmic equations were examined and correction factors for these errors are presented. i i Estimates of component biomass were obtained using allometric equations, the stand-table and height-diameter data from eighty-eight of 100-year-old lodgepole pine trees covering a range of site and stand density condition. The combined variable of stand basal area times mean stand height (BA'H) was closely associated with most component dry-weights per acre. Although total tree and component biomass per acre were positively correlated with basal area per acre they were inversely related to number of stems per acre on a given site. Fresh- and dry-weight estimates, by component, were obtained from two paired, young lodgepole pine and Populus stands grown in west central Alberta. Equations for the estimation of the component dry-weights of young lodgepole pine trees are presented. For the sites examined, the above-ground total stand crop of the young lodgepole pine stands was substantially higher than that of similarly aged Populus stands grown on similar sites. Some possible reasons for these d i f f -erences in productivity are discussed. Within and between tree variations in radial, cross-sectional area, and volume growth were examined for twenty, 100-year-old lodge-pole pine trees. Ten trees were sampled from both a thinned and an unthinned stand. Although volume growth was found to be highly cor-related with the amount of foliage, volume growth efficiency (growth per unit of foliage) was not related to tree size. Equations for relating the amount of growth at any point in the tree to the needle biomass at or above that point in the tree are presented. A large amount of within tree variation in growth was not accounted for by these relation-ships. Thinning did not appear to affect the pattern of growth of the trees examined. Suggestions for further research on the growth patterns i i i as related to foliage biomass are presented. Ten pulp sample trees, two from each of the 4-, 6-, 8-, 10-, and 12-inch diameter classes, were collected from two average density, 100-year-old stands of lodgepole pine grown in southwestern Alberta. The oven-dry, bark-free weights of the merchantable stem (4.0-inch top), non-merchantable top (4.0- to 1.0-inch top), branch (> 1.0 inch diameter), and root-stump (> 1.0 inch diameter) components were measured for each tree. The relationships between the quantity of these components, expressed as a percentage of the oven-dry, bark-free fu l l bole, and tree size are presented. The relationships between tree size, and the yield and quality of kraft pulp produced from each component are examined. A significant positive relationship was found between tree size and the unscreened yield of pulp from the f u l l bole component only. No significant relationships were found between tree size or growth rate and pulp quality for any of the components. Variations in the yield and quality of kraft pulp at different locations within a single tree are examined and discussed. Because the yield and quality of pulp from the non-merchantable top was found to be relatively high and because quantity of this component per acre may be substantial in some stands, immediate consideration should be given to its utilization. Although the yield and quality of pulp from the root-stump system was found to be high, utilization of this component in the near future is doubtful because of the technical problems associated with extraction, cleaning, transporation, and processing. Similarly, utilization of branches in the near future is doubtful because of the low yield and quality of pulp from branchwood and because of the processing problems associated with its utilization. iv Nine highly suppressed, 100-year-old lodgepole pine trees from a high density stand in southwestern Alberta were collected for pulping studies. Kraft pulp yield and quality data are presented for the bole and root-stump components. The results demonstrate that the utilization of these small diameter trees for the manufacture of pulp depends upon economic harvesting and processing and not pulp yield and quality. V ACKNOWLEDGEMENTS The writer wishes to express his sincere thanks to Dr. D. D. Munro for his guidance, advice, and encouragement. The writer is also greatly indebted to Dr. A. Kozak, Dr. J . H. G. Smith and Mr. S. M. Smith, of the Faculty of Forestry, and Dr. J. L. Keays, of the Western Forest Products Laboratory, for their critical review, advice and encouragement. The assistance of Dr. A. Kozak and Miss L. Cowdell, in programming, plotting and analysing the data, and the assistance of Dr. J . H. G. Smith, in the measurement of earlywood and latewood, are gratefully acknowledged. The biomass and complete-tree utilization data used in this thesis were collected between 1966 and 1972 by the writer while he was employed as a Research Officer with the Canadian Forestry Service. The writer would like to thank the Canada Department of the Environment, Canadian Forestry Service, for making the data used in this thesis available. Sincere thanks are due, to Mr. Stan Lux, for his assis-tance in the field and laboratory work, and for the draughting; and to other members of the Northern Forest Research Centre's staff, for assisting in the preparation and typing of the manuscript. Special thanks are also due, to Dr. J . L. Keays, Dr. J . V. Hatton and the technical staff of the Pulping Section of the Western Forest Products Laboratory for their advice and assistance in the complete-tree utilization phase of this thesis. Attendance at the University was facilitated by financial assistance from the Canadian Forestry Service, and by financial assistance from the Faculty of Forestry, University of British Columbia, in the form of fellowships and assistantships. Finally, I wish to express my special appreciation to my wife Beverley, and to my sons David and Ian for their patience, understanding, encouragement and sacrifice throughout this study. v i i TABLE OF CONTENTS Page ABSTRACT i ACKNOWLEDGEMENTS v TABLE OF CONTENTS v i i LIST OF TABLES xi LIST OF FIGURES xv CHAPTER I INTRODUCTION 1 CHAPTER II COMPONENT DRY-WEIGHTS OF 100-YEAR-OLD LODGEPOLE PINE TREES AND STANDS 4 Introduction 4 Methods and Materials 5 Data collection 5 Analysis 11 Results and Discussion 16 Component dry-weights of 100-year-old lodgepole pine trees 16 Component dry-weights of 100-year-old lodgepole pine stands 28 Conclusions 36 CHAPTER III DRY-MATTER COMPARISONS BETWEEN YOUNG LODGEPOLE PINE AND POPULUS STANDS 39 Introduction 39 Methods and Materials 40 Data collection 40 Analysis 43 Results and Discussion 44 TABLE OF CONTENTS (Continued) Page Dry-weight relationships 45 Fresh-weight relationships 47 Above-ground organic matter production of paired lodgepole pine and Populus stands of similar ages on similar sites 51 Conclusions 54 CHAPTER IV VARIATIONS IN STEM GROWTH AS RELATED TO SEVERAL CROWN CHARACTERISTICS OF 100-YEAR-OLD LODGEPOLE PINE TREES 56 Introduction 56 Methods and Materials 58 Data collection 58 Analysis 60 Results and Discussion. 61 Conclusions 71 CHAPTER V COMPLETE-TREE UTILIZATION OF AVERAGE STAND DENSITY 100-YEAR-OLD LODGEPOLE PINE... 74 Introduction 74 Methods and Materials. 76 Data collection 76 Analysis 84 Results and Discussion 87 The quantity of each component potentially available from the complete-tree utilization of 100-year-old lodgepole pine trees 87 The quality of pulp obtained from the complete-tree utilization of 100-year-old lodgepole pine trees 91 TABLE OF CONTENTS (Continued) Page V a r i a t i o n among trees 91 V a r i a t i o n w i t h i n trees 97 Conclusions 102 CHAPTER VI COMPLETE-TREE UTILIZATION OF HIGH STAND DENSITY 100-YEAR-OLD LODGEPOLE PINE..... 107 Introduction 107 Methods and Mat e r i a l s 108 Data c o l l e c t i o n 108 Preparation and t e s t i n g of paper from the bole component 109 Preparation and t e s t i n g of paper from the root-stump component.. I l l An a l y s i s . . 113 Results and Discussion 113 Conclusions 115 CHAPTER VII SUMMARY AND SUGGESTIONS FOR FURTHER RESEARCH 117 REFERENCES CITED 121 APPENDICES: I Lesser vegetation present i n Stands 1 and 2 132 I I - l Dry t o t a l tree weight - DBH allometry of mature white spruce 133 II-2 Dry above-ground weight - DBH allometry of mature white spruce 133 II-3 Dry stem weight - basal area r e l a t i o n s h i p of mature white spruce 133 II-4 Dry merchantable stem weight - bas a l area r e l a t i o n s h i p of mature white spruce 133 TABLE OF CONTENTS (Continued) Page II-5 Dry needle weight - DBH allometry of mature white spruce 134 II-6 Dry branch weight - DBH allometry of mature white spruce 134 II-7 Dry root-stump weight - basal area relationship of mature white spruce 134 III Radial growth and foliage weight distribution diagrams of twenty, 100-year-old lodgepole pine trees 135 xi LIST OF TABLES Table Page 1 Characteristics of Stands 1 and 2 in 1938 before and after thinning... 6 2 Characteristics of the three stands of 100-year-old lodgepole pine . 6 3 Statistical characteristics of the independent variables for 100-year-old lodgepole pine trees 16 4 Statistical characteristics of the dependent variables for 100-year-old lodgepole pine trees 17 5 Simple correlation coefficients (r) between tree and component dry-weights and several tree characteristics for 100-year-old lodgepole pine trees 18 6 Dry-weight allometric relationships for 100-year-old lodgepole pine trees 20 7 Dry weight of tree components and total standing crop for three stands of 100-year-old lodgepole pine 27 8 Statistical characteristics of the independent variables from eighty-eight stands of 100-year-old lodgepole pine 28 9 Statistical characteristics of the dependent variables from eighty-eight stands of 100-year-old lodgepole pine... 29 10 Simple correlation coefficients (r) between several stand characteristics and stand component dry-weights for eighty-eight stands of 100-year-old lodgepole pine.... 29 11 Regression statistics for evaluating the influences of site quality, number of stems per acre and basal area per acre on the total standing crop of 100-year-old lodgepole pine stands 35 12 Characteristics of the four lodgepole pine and Populus stands 41 13 Statistical characteristics of the independent variables from twenty, young lodgepole pine trees 45 14 Statistical characteristics of the dry-weight dependent variables from twenty, young lodgepole pine trees 45 x i i LIST OF TABLES (Continued) Table Page 15 Simple correlation coefficients (r) between component dry-weights and several tree characteristics of twenty, young lodgepole pine trees 46 16 Regression equations, and regression coefficient significance tests (Fx^), derived from twenty, young lodgepole pine trees, used to estimate sample plot living component dry-weights 47 17 Statistical characteristics of the fresh-weight dependent variables from twenty, young lodgepole pine trees 49 18 Simple correlation coefficients (r) between component fresh-weights and several tree characteristics of twenty, young lodgepole pine trees 49 19 A comparison of measured fresh weights (lb./ac.) of paired lodgepole pine and Populus stands of similar ages on similar sites 52 20 A comparison of estimated above-ground dry weight (lb./ac.) of living components in paired lodgepole pine and Populus stands of similar ages on similar sites 51 21 Proportion of total above-ground dry-weight by components of major species in study areas 54 22 Some characteristics of the twenty, 100-year-old lodgepole pine sample trees 62 23 Simple correlation coefficients (r) between several tree characteristics of the twenty, 100-year-old lodgepole pine sample trees 63 24 Analyses of variance of radial, cross-sectional area, and volume growth measured at 5.positions within the crown and 5 positions within the clear-bole of ten, 100-year-old lodgepole pine trees grown in a thinned stand and ten, 100-year-old lodgepole pine trees grown in an unthinned stand 66 i x i i i LIST OF TABLES (Continued) Table Page 25 Simple correlation coefficients (r) between 358 measurements of radial, cross-sectional area and section volume growth, and several tree and section characteristics . 67 26 Simple correlation coefficients (r) between 245 measurements of radial, cross-sectional area and section volume growth in crown-formed wood, and several tree and section characteristics... 69 27 Comparison of relative values of kraft pulp from conifer tree components expressed in terms of the bole 75 28 Characteristics of the selected 100-year-old lodgepole pine trees... 79 29 Kraft cooking conditions used for the exploratory cooks 81 30 Kraft cooking conditions used in study to obtain unbleached pulps with permanganate numbers of approximately 20 from the three components of 100-year-old, forest-grown lodgepole pine trees 81 31 Bark-free, oven-dry weights of the 100-year-old lodgepole pine tree components as a percentage of f u l l boles 87 32 Average number of stems per acre in diameter classes less than 9.0 inches dbhob from eighty-eight stands of 100-year-old lodgepole pine 90 33 Component pulp yield data from kraft pulping for ten, 100-year-old lodgepole pine trees 92 34 Component pulp quality at 300 ml CSF from kraft pulping for ten, 100-year-old lodgepole pine trees... 93 35 Simple correlation coefficients (r) between tree size (D) and component unscreened pulp yield (at permanganate number 20) and component unbleached pulp quality (at 300 ml CSF) from kraft pulping ten, 100-year-old lodgepole pine trees 94 36 Analysis of variance of duplicate determinations of f u l l bole pulp yield for two trees in each of five diameter classes of 100-year-old lodgepole pine trees 95 xiv LIST OF TABLES (Continued) Table Page 37 General differences in the wood and pulp characteristics of non-merchantable tops compared with merchantable boles for coniferous species 98 38 A comparison of the yield and quality of unbleached kraft pulps from the merchantable bole, non-merchantable top, and f u l l bole of a 100-year-old lodgepole pine tree 101 39 Characteristics of the nine suppressed, 100-year-old lodgepole pine trees used in this pulp study 109 40 Kraft cooking conditions for the bole chip mixture of nine suppressed, 100-year-old lodgepole pine trees 110 41 Kraft cooking conditions for the root-stump chip mixture of nine suppressed, 100-year-bld lodgepole pine trees I l l 42 A comparison between unscreened pulp yield data of suppressed pine trees, more normally grown pine trees and the reference standard 114 43 A comparison between the mean unbleached pulp quality of the suppressed pine trees, more normally grown pine trees and the reference standard 115 XV LIST OF FIGURES Figure Page 1 The relationship between moisture content and height in the tree 8 2 The relationship between specific gravity and height in the tree 8 3 The relationship between bias correction factors and standard errors of estimate (a) of logarithmic equations 15 4 Total tree weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine 21 5 Total above-ground component weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine 21 6 Stem weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine 21 7 Merchantable stem weight - D2H relationship of 'average stand density' 100-year-old lodgepole pine 21 8 Stem bark weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine 22 9 Needle weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine 22 10 Branch weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine 22 11 Stump plus root weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine 22 12 Root weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine 23 13 Total tree weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine 24 14 Total above-ground component weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine 24 15 Stem weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine 24 xyi LIST OF FIGURES (Continued) Figure Page 16 Needle weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine 24 17 Branch weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine 25 18 Stump plus root weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine 25 19 The relationship between estimated dry needle weight per acre and stand basal area times mean stand height for stands of 100-year-old lodgepole pine 31 20 The relationship between estimated dry branch weight per acre and stand basal area times mean stand height for stands of 100-year-old lodgepole pine 31 21 The relationship between estimated dry stem weight per acre and stand basal area times mean stand height for stands of 100-year-old lodgepole pine 31 22 The relationship between estimated dry above-ground weight per acre and stand basal area times mean stand height for stands of 100-year-old lodgepole pine 31 23 The relationship between estimated dry root plus stump weight per acre and stand basal area times mean stand height for stands of 100-year-old lodgepole pine 32 24 The relationship between estimated dry total tree weight per acre and stand basal area times mean stand height for stands of 100-year-old lodgepole pine 32 25 The relationship between estimated dry total tree weight per acre and number of stems per acre for stands of 100-year-old lodgepole pine 33 26 The relationship between stem dry-weight and tree diameter of young lodgepole pine trees 48 xvii LIST OF FIGURES (Continued) Figure Page 27 The relationship between branch dry-weight and tree diameter of young lodgepole pine trees 48 28 The relationship between needle dry-weight and tree diameter of young lodgepole pine trees 48 29 The relationship between above-ground dry-weight and tree diameter of young lodgepole pine trees 48 30 The relationship between stem fresh-weight and D2H of young lodgepole pine trees 50 31 The relationship between crown fresh-weight and D2H of young lodgepole pine trees 50 32 The relationship between above-ground fresh-weight and D2H of young lodgepole pine trees 50 33 The relationship between needle dry-weight and tree diameter for the twenty, 100-year-old lodgepole pine trees 64 34 The relationship between past 5-year volume growth and present needle dry-weight of twenty, 100-year-old lodgepole pine trees 64 35 The relationship between volume growth efficiency (cu.ft./lb. dry needles) and tree diameter of twenty, 100-year-old lodgepole pine trees 64 36 The relationship between unbleached pulp permanganate number and effective alkali for three components of 100-year-old lodgepole pine 82 37 Relationship between unscreened yield and permanganate number for kraft pulp from three components of 100-year-old lodgepole pine trees... 86 x v i i i LIST OF FIGURES (Continued) Figure Page 38 The relationship between tree size and oven-dry, bark-free component weight expressed as a percentage of the oven-dry, bark-free f u l l bole weight for 100-year-old lodgepole pine trees 89 39 The relationship between total unscreened pulp yield and tree size 96 40 Unscreened yield (at 20 permanganate number) for kraft pulp from several locations within a 100-year-old lodgepole pine tree 99 41 Burst Factor (at 300 ml CSF) for unbleached kraft pulp from several locations within a 100-year-old lodgepole pine tree 100 42 Tear factor (at 300 ml CSF) for unbleached kraft pulp from several locations within a 100-year-old lodgepole pine tree 100 43 Breaking length (at 300 ml CSF) for unbleached kraft pulp from several locations within a 100-year-old lodgepole pine tree 100 44 Bulk factor (at 300 ml CSF) for unbleached kraft pulp from several locations within a 100-year-old lodgepole pine tree 100 CHAPTER I INTRODUCTION As man becomes more conscious of his environment he demands a greater knowledge of the biological and physical factors, including the flow of organic matter, energy, nutrients, and moisture which affect natural ecosystems. Although foresters have traditionally measured the forest ecosystem in terms of. yield to man (i.e., volume per unit area to a certain merchantability limit set by current technological and economic conditions) such measures are of l i t t l e direct value in assessing the total productivity and dynamics of the forest ecosystem. It is for these reasons that foresters are becoming increasingly interested in determining forest dry-matter production and biomass. The measurement of forest biomass dates back to the classic studies by Burger (1929), Dengler (1937), Kittridge (1944), and MartMoller (1947). During the past two decades increased attention has been focused upon the functioning and productivity of forest eco-systems particularly with the advent of the International Biological Programme (IBP). This increased attention is the result of greater interest in quantitative ecology (Ovington, 1957, 1962; Baskerville, 1965a, 1966; Tadaki, 1966; Whittaker, 1966; Rodin and Bazilevic, 1965, 1966, 1968; Kira and Shidei, 1967; Satoo, 1967), forest fertilization and nutrient flow (Ovington and Madgwick, 1959; Orman and Will, 1960; Will, 1964; Young et a l . , 1964; Rennie, 1955; 2 Bazilevic and Rodin, 1966, Marchenko, 1967; Duvigneaud and Denaeyer-De Smet, 1967; Woodwell and Whittaker, 1967; Bormann et a l . , 1967; Cole et a l . , 1967; Young and Carpenter, 1967; White et a l . , 1971), forest harvesting (Keen, 1963; Adamovich, 1970), and complete-tree utilization (Young and Chase, 1965; Keays,1968, 1971a). Comprehensive reviews of past biomass studies have been presented by Ovington (1962), Baskerville (1965a), Tadaki (1966), Johnstone (1967), Kira and Shidei (1967), and Keays (1971b,c,d,e,f). Biomass and net annual primary productivity data have been summarized by Ovington (1962), Rodin and Bazilevic (1965), Tadaki (1966), and Art and Marks (1971). Plants, through the process of photosynthesis (and because of the unique properties of the carbon atom), transform radiant energy into the form of utilizable chemical energy. The process of photo-synthesis, wherein carbohydrates are formed from carbon dioxide and water, is the primary source of organic matter and potential energy upon which a l l l i f e (with the exception of some bacteria) is dependent. The productivity and efficienty of any plant organism within the ecosystem depends upon the morphology, anatomy, and physiology of that organism. The productivity and efficiency of the ecosystem varies with the aggregate productivity and efficiency of the individual organisms, the interaction between organisms, reaction to edaphic, climatic and physiographic influences, and the influence of man. However, as Madgwick (1970) pointed out, l i t t l e is known of the distribution of photosynthate within forest stands and how to successfully manipulate this distribution to maximize harvests. Because of the laborious, destructive, and costly nature of the measurements necessary in studying dry-matter production and 3 complete-tree utilization i t may be advantageous to limit studies to an intensive examination of a small number of individuals in a small number of locations. In so doing, data could be obtained pertaining to studies of biomass, complete-tree utilization and tree growth, which would otherwise require three separate studies, resulting in lower field costs and greater meaningfulness through complete compatability among studies. Although the results obtained from such an integrated study suffer from lack of generality this could be rectified by continued study. The German Soiling Project (Ellenberg, 1971) is the best documented, multi-disciplinary ecosystem study available to date. The objective of this thesis is to present the results of studies of dry-matter production, tree growth, and complete-tree utilization of 100-year-old lodgepole pine (Pinus  contorta Dougl. var. la t i f o l i a Engelm.) based on data obtained from the intensive examination of a relatively few forest-grown trees from three stands in southwestern Alberta. As a first approximation, the results thus obtained are used to compare the dry-matter production of similarly aged lodgepole pine stands grown in other regions of Alberta. In addition, comparisons are made between the above-ground dry-matter production of young lodgepole pine and Populus stands i n Alberta. 4 CHAPTER I I COMPONENT DRY-WEIGHTS OF 100-YEAR-OLD LODGEPOLE PINE TREES AND STANDS Introduction In Canada biomass r e s u l t s have been reported f o r Picea  mariana ( M i l l . ) BSP (Weetinan and Harland, 1964), Abies balsamea (L.) M i l l . ( B a s k e r v i l l e , 1965a,b, 1966; Honer, 1971), Pinus contorta Dougl. var. l a t i f o l i a Engelm. ( K i i l , 1965; Muraro, 1966; Johnstone, 1967, 1970a,b, 1971), Pinus resinosa A i t . ( S t i e l l , 1966), Populus  tremuloides Michx. ( B e l l a and J a r v i s , 1967; B e l l a , 1968, 1970; Peterson e t a l . , 1970; P o l l a r d , 1972), Picea glauca (Moench) Voss (Keays and Hatton, 1971b), Pinus banksiana Lamb. (Hegyi, 1972), and Tsuga he t e r o p h y l l a (Raf.) Sarg., Pseudotsuga menziesii (Mirb.) Franco, and Thuja p l i c a t a Donn (Osborn, 1968; Kurucz, 1969; P a i l l e , 1970; Keays and Hatton, 1971a; Smith, 1971; McGreavy, 1972). The purpose of t h i s chapter i s to examine the dry weight and d i s t r i b u t i o n of the various components of 100-year-old lodgepole pine trees, to develop a l l o m e t r i c r e l a t i o n s h i p s which e x i s t between the dry weights of the components and seve r a l e a s i l y measured tree c h a r a c t e r i s t i c s , and to compare the t o t a l standing crop of 100-year-o l d lodgepole pine stands growing under d i f f e r e n t s i t e and stand density conditions i n A l b e r t a . 5 Methods and Materials Data collection The data for this study were gathered from sample plots located in each of three stands on the Kananaskis Forest Research Station (North Latitude 51°06', West Longitude 115°04') in the SA 1 Section of the Subalpine Forest Region (Rowe, 1959). The three stands were on a well-drained, gently sloping site, of medium site quality, at an elevation of about 4,600 feet. The soil is a calcareous grey podzol. The species composition of the stands was predominantly lodgepole pine with some white spruce (Picea glauca Moench) Voss var. albertiana (S. Brown) Sarg.) present. Appendix I lists some of the most prominent lesser vegetation present in the plots. The trees were a l l about 100 years old. Before 1938 Stands 1 and 2 were very similar in terms of basal area per acre, total volume per acre, and average stand diameter (Table 1). In 1938, when the trees were about 70 years old, Stand 2 was subjected to a crown thinning, which removed about 39 percent of the number of trees, 31 percent of the basal area, and 30 percent of the volume, principally from the smaller diameter classes (Table 1). Unfortunately, the weight of the material removed by thinning is not known. No treatment was applied to Stand 3 and the condition of this stand in 1938 is unknown. When the present study was carried out most of the trees in Stand 3 were very suppressed and many were infected by dwarf mistletoe (Arceuthobium americana Nutt. ex Engelm.) and atropellis canker (Atropellis piniphila (Weir) Lohman and Cash). 6 Table 1. Characteristics of Stands 1 and 2 in 1938 before and after thinning. Stand 1 Stand 2 Characteristics Before After Thinning Thinning Living stems per acre Basal area per acre (sq.ft.) Volume per acre (cu. ft.) Mean dbh (in.) 3,005 2,003 1,220 195.6 187.6 128.6 3,905 4,149 2,910 3.5 3.6 4.4 Because measurement of a l l of the trees was impractical due to the technical problems of handling and weighing the trees, i t was decided to use sampling with regression to estimate stand biomass. One square tenth-acre plot was established in each of Stands 1 and 2, and a rectangular twentieth-acre plot was located in Stand 3. The characteristics of the stands, based on plot measurements are presented in Table 2. Table 2. Characteristics of the three stands of 100-year-old lodgepole pine. Characteristic Stand 1 Stand 2 Stand 3 (Thinned) Living stems per acre 1,020.0 290.0 4,960.0 Basal area per acre (sq.ft.) 227.7 152.0 156.5 Volume per acre (cu.ft.) 6,356.0 5,107.0 2,594.0 Mean dbh (in.) 6.5 9.8 2.2 Mean height (ft.) 54.7 66.5 18.7 The trees were felled at 1 foot above the ground. Diameter at breast height to the nearest 1/10 inch, total tree height to the nearest 1/10 foot, crown width (the average of two measurements taken at right angles at the widest part of the crown) to the near-7 est 1/10 foot, and live crown length (the length from the tip to the lowest whorl of live branches) to the nearest 1/10 foot were measured. A dial scale with a capacity of 500 pounds was used to weigh the trees. The total tree above stump height (including branches and needles), the total stem above stump height (total tree less branches and needles), and the merchantable stem (4.0-inch top diameter outside bark) were weighed to the nearest 1.0 pound. In addition, stem maps showing the exact location of each tree and its crown were prepared for each sample plot. Discs, each about one inch thick, were sawn from the stem at stump height, breast height, and at eight-foot intervals above stump height to the top of the tree. The diameter outside bark to the nearest 1/10 inch was measured with a diameter tape and recorded for each disc. The discs were placed in sealed polyethylene bags to minimize moisture loss during transport to the laboratory ( a maximum time period of 3 hours). In the laboratory, the bark was separated from each disc and the diameter inside bark of each disc was measured. The wood and bark of each disc were weighed separately and placed in a drying oven at 105°C until a constant oven-dry weight was determined. Because of the within tree variations in moisture content (Figure 1), an arith-metic average moisture content was not deemed representative of the entire tree. Therefore, an average moisture content for each entire tree was determined by weighing the average moisture content of two consecutive discs by the volume (calculated by Smalian's formula) of the section between the two discs. A similar method was used to calculate a weighted average bark moisture content. These weighted Figure 2. THE RELATIONSHIP BETWEEN SPECIFIC GRAVITY A N D HEIGHT IN THE TREE.' 0600 Spgr» 0.497 - 0.000854 H (ft) Sy.x • 0.039 r2« O.I37**n » 713. 20 30 40 50 H e i g h t in Tree [ F t . a b o v e G r o u n d ] 60 70 S o u r c e • J o h n s t o n e [ 1 9 7 0 a ] 9 average moisture contents were used to determine stem wood, and stem bark, dry-weights. The actual fresh volume inside and outside bark was determined from Reineke charts. Bark dry-weight was estimated by multiplying bark volume times lodgepole pine bark density (Smith and Kozak, 1967). A sector, containing a line of mean radius, was sawn from each oven-dry disc. Each sector was redried for 24 hours at 105°C, weighed to the nearest 1/100 gram, and covered with a very thin coating of paraffin. The specific gravity of each sector was deter-mined by Method B-II of the ASTM Designation D2395-65T (3) procedure (American Society for Testing and Material, 1967). The only devia-tions from the ASTM procedure were as follows: 1. some of the sectors were not free from knots and other defects, and 2. no attempt was made to extract the pitch from the sectors. Because specific gravity varies with height within a tree (Figure 2), a weighting procedure, similar to that described above for determining a weighted average moisture content, was used to calculate a weighted average specific gravity for each tree. After the entire tree above stump height was weighed the needle-bearing twigs were clipped from the larger branch parts, and put into burlap sacks. The burlap sacks were placed in a drying shed in which the temperature was maintained at about 85°C. Repeated tests revealed that a two-week drying period was required to ensure that the foliage had reached a constant oven-dry moisture content. The dried needles were removed from the twigs by hand and the cleaned 10 needles (without fascicles) were weighed. Needle moisture-content data ( K i i l , 1968) were used to estimate the fresh needle weight of each tree. The weight of fresh branches of each tree was obtained by subtracting the estimated weight of fresh needles from the measured fresh weight of crown materials (needles plus branches). The dry branch weight of the tree was then estimated by reducing the calculated fresh branch weight by the moisture content of branch wood ( K i i l , 1968). A D-8 Caterpillar tractor was used to uproot the root-stump components of each tree. The roots were washed free of soil particles and weighed after the surface had dried. Discs were cut from the root and stump components so that dry-weight calculations could be made. Although some root materials (e.g. small roots and root hairs) were lost in this method of extraction, the losses, in terms of their dry-weight, were small. Data from 88 sample plots, ranging in size from 1/20 to 1/2 acre, established in essentially pure, 100-year-old (91-110 years) lodgepole pine stands (stands in which the basal area of species other than pine was less than 25 percent of the total pine basal area) were chosen to represent a wide variety of age, site, and stand density conditions. For each sample plot the following data was obtained: 1. A stand table and height-diameter curve. 2. Total age. 3. An estimate of site index (Kl) based on the conventional method of. using unadjusted dominant and codominant height and index age 70 (Kirby, 1968). 4. A second estimate of site index (AI) wherein observed dominant height was expressed as a percentage of dominant 11 height as estimated from the following equation: Log Dom. Ht. = 2.7378-9.6252 (l/Age)-0.28576 (Log No. 10 10 of Stems). Because the data used to derive the above equation were collected from a wide range of age, site, and stand density conditions the equation was deemed to represent an average for the species in Alberta. Using the allometric equations developed from Stand 1 and 2, and the stand-table and height-diameter data from each plot i t was possible to estimate the total standing crop of lodgepole pine stands for a wide range of age, stand density and site types. Analysis Because the thinning did not affect the weight or distribution of the trees' components, the data from Stands 1 and 2 (hereafter referred to as 'average stand density') were pooled and analyzed together. Because the heavy suppression experienced in Stand 3 did affect the distribution and weight of the various components, the data from this stand (hereafter referred to as'high stand density') were analyzed separately. A l l of the data were analyzed by multiple regression techniques with the computer program described by Kozak and Smith (1965). Tree component dry-weights (Table 4), in pounds, were used as dependent variables with the tree characteristics presented in Table 3 as the independent variables. The following were used as dependent variables in the regression analyses of the tree and component weights: a) Total tree dry-weight (Y ): the dry weight of a l l 12 components including needles, branches, cones, bole wood, bark and the root-stump component. b) Total above-ground dry-weight (Y ): the dry weight of 2 a l l components above a 1.0-foot stump. c) Total stem dry-weights (Y ) : the dry weight of the stem, 3 including bark, from the top of the tree to a 1.0-foot stump. d) Merchantable stem dry-weight (Y ): the dry weight of that portion of the stem, including bark, from a 1.0-foot stump to a 4.0-inch top. e) Stem bark dry-weight (Y ): the dry weight of stem bark above a 1.0-foot stump. f) Needle dry-weight (Y ): the dry weight of needles. 6 g) Branch dry-weight (Y ): the dry weight of living branches. 7 h) Stump plus root dry-weight (Y ): the dry weight of a 1.0-8 foot stump plus roots. i) Root dry-weight (Y ): the dry weight of that portion 9 of the tree below ground level. The regression equations presented in the following section for tree and tree component weights with the exception of merchantable stem dry-weight, are of a logarithmic transformation (allometric function) form. Logarithmic transformations were used to facilitate the applic-ation of a linear model and to permit comparisons with similar work reported in the literature which most frequently use this transformation. In the following results the standard error of estimate is expressed both in absolute units and as a percentage of the mean, the latter being in parentheses. Because the standard error of estimate 13 determined from the r e s i d u a l mean square of a logarithmic equation cannot be transformed back to an arithmetic scale without bi a s , the standard errors of estimate were ca l c u l a t e d by the following formula: where: Y = observed value of the dependent v a r i a b l e 5 Y = estimated value of the dependent v a r i a b l e n = number of observations S i m i l a r l y , an estimated c o e f f i c i e n t of determination ( r 2 ) was obtained by the following formula: 2 2 R or r = SStotal-SSresid. SStotal where: SStotal = sum of squares of untransformed Y SSresid.= I (Y-Y) 2 Because the squared deviations of logarithmic values rather than arithmetic values are minimized, the use of logarithmic equations, derived by the l e a s t squares methods, r e s u l t s i n a systematic under-estimate. A method of overcoming t h i s problem was suggested by Meyer (1944) based on the following formula: 2 C F = 101.1513-<* where: CF = c o r r e c t i o n factor a = standard er r o r of estimate of a logarithmic equation i n terms of logarithms ( i . e . , as determined from the r e s i d u a l mean square) Figure 3 presents t h i s c o r r e c t i o n f a c t o r (CF) g r a p h i c a l l y . Parameter estimates from a logarithmic equation need merely be m u l t i p l i e d by the appropriate c o r r e c t i o n f a c t o r f or that equation to obtain unbiased 14 estimates. As can be seen from Figure 3 the application of the correction factor becomes critical when the o* exceeds 0.10. In addition, aggregate deviations (AD) were calculated for each equation by determining the difference between the sum of observed A. weights (IY) and the sum of estimated weights (EY) expressed as a decimal fraction of the latter using the following formula: EY 1'5 Figure 3. THE RELATIONSHIP BETWEEN BIAS CORRECTION FACTORS AND STANDARD ERRORS OF ESTIMATE (a) OF LOGARITHMIC EQUATIONS. 20 r 16 Results and Discussion Component dry-weights of 100-year-old lodgepole pine trees Table 3 presents the means, standard deviations, and minimum and maximum values of the independent v a r i a b l e s used i n the analyses. Table 3. S t a t i s t i c a l c h a r a c t e r i s t i c s of the independent v a r i a b l e s f o r 100-year-old lodgepole pine trees. Independent Mean Stand Minimum Maximum va r i a b l e s dev. value value Average stand density (Stands 1 and 2 pooled) - 85 observations Diameter (D) (in.) 7.1 2.1 4.0 13.4 Height (H) ( f t . ) 60.4 7.5 45.0 81.7 Crown length (CL) ( f t . ) 21.6 10.3 8.0 54.0 Crown width (CW) ( f t . ) 5.4 2.0 2.5 14.7 Stump age (AGE) (yrs.) 92.4 4.9 75.0 100.0 Height to l i v e crown (HLC) (ft.)38.8 6.6 20.1 50.8 Diameter squared (D 2) ( i n . 2 ) 54.3 32.7 16.0 179.6 Diameter squared times height (D2H) ( i n . 2 • f t . ) 3,498.4 2 ,581.0 760.0 13,772.2 High stand density (Stand 3) - 221 observations Diameter (D) (in.) 2.2 1.0 0.6 6.6 Height (H) ( f t . ) 18.8 6.7 5.7 41.0 Crown length (CL) ( f t . ) 9.4 4.8 1.6 27.0 Crown width (CW) ( f t . ) 2.5 1.0 0.5 6.5 Height to l i v e crown (HLC) ( f t . ) 9.3 2.8 3.7 18.2 Diameter squared (D 2) ( i n . 2) 5.8 5.7 0.4 43.6 Diameter squared times height (D 2H) ( i n . 2 • f t . ) 141.9 205.3 2.1 1,781.6 The means, standard deviations, and maximum and minimum values of the dependent v a r i a b l e s are presented i n Table 4. 17 Table 4. S t a t i s t i c a l c h a r a c t e r i s t i c s of the dependent v a r i a b l e s for 100-year-old lodgepole pine t r e e s . Independent Mean Stand Minimum Maximum v a r i a b l e dev. value value Average stand density (Stands 1 and 2 pooled) - 85 observations T o t a l tree dry-weight (Yi) (lb, .) 372.7 280 .1 92.3 1,530.3 T o t a l above-ground dry-weight (Y2) (lb .) 311.4 226 .2 64.3 1,238.1 T o t a l stem dry-weight (Y 3) (lb .) 268.9 179 .8 58.9 944.3 Merchantable stem dry-weight (Y 4) (lb .) 237.3 186 .4 0.0 930.0 Stem bark dry-weight <*5> (lb • ) 31.2 29 .7 5.3 170.8 Needle dry-weight (Y 5) (lb • ) 14.0 11 .7 1.0 61.2 Branch dry-weight (Y 7) (lb .) 21.8 30.8 0.4 194.9 Stump plus root dry-weight (Y 8) (lb .) 57.4 49 .8 13.2 292.2 Root dry-weight (Y 9) (lb .) 51.5 48 .1 9.1 283.5 High stand density (Stand 3) - 221 observations T o t a l tree dry-weight (Yi) (lb .) 20.8 27 .2 0.8 214.8 T o t a l above-ground dry-weight (Y 2) (lb • ) 16.8 22 .2 0.6 173.8 T o t a l stem dry-weight (Y 3) (lb .) 12.5 16 .0 0.4 125.4 Needle dry-weight (Y 6) (lb .) 1.4 2 .2 0.02 16.1 Branch dry-weight (Y 7) (lb .) 2.9 4 .3 0.04 32.3 Stump plus root dry-weight (Y 8) (lb .) 4.0 5 .1 0.2 41.0 72 observations were used f o r root plus stump weight, root weight, and t o t a l tree weight, t h e r e f o r e , the dependent v a r i a b l e s are not a d d i t i v e . Simple c o r r e l a t i o n c o e f f i c i e n t s (r) between the untransformed dependent and independent v a r i a b l e s are presented i n Table 5. Table 5. Simple correlation coefficients (r) between tree and component dry-weights and several tree characteristics for 100-year-old lodgepole pine trees. Independent Dependent variables variables1 Y2 Y3 Y«+ Y5 Y6 Y7 Y8 Y9 Average stand density (Stands 1 and 2 pooled) - 85 observations3 D 0.961 0.966 0.975 0.979 . 0.870 0.921 0.814 0.914 0.899 H 0.882 0.894 0.912 0.917 0.852 0.841 0.713 0.825 0.809 CL 0.783 0.796 0.798 0.800 0.833 0.782 0.694 0.753 0.744 cw 0.894 0.888 0.873 0.879 0.831 0.883 0.867 0.885 0.879 AGE 0.098 0.155 0.173 0.180 0.074 0.185 0.034 0.060 0.055 HLC -0.235 -0.222 -0.204 -0.202 -0.328 -0.262 -0.251 -0.251 -0.256 D2 0.989 0.989 0.989 0.990 0.900 0.940 0.881 0.960 0.950 D2H 0.993 0.992 0.989 0.989 0.922 0.940 0.902 0.972 0.965 High stand density (Stand 3) - 221 observations D 0.922 0.921 0.924 0.873 0.873 0.897 H 0.871 0.871 0.879 0.812 0.812 0.847 CL 0.859 0.858 0.853 0.832 0.832 0.841 CW 0.725 0.727 0.721 0.709 0.709 0.697 HLC 0.606 0.608 0.656 0.513 0.513 0.581 D2 0.983 0.982 0.983 0.936 0.936 0.959 D2H 0.992 0.990 0.992 0.943 0.943 0.970 p.05 signif. level ^ y Q « 0.232 r 83= 0.216 r - 0.137 1 * 9 y See Table 3 for definition of abbreviations. 2 See Table 4 for definition of abbreviations. 3 72 observations were used for root plus stump, root, and total tree dry-weights. 19 As can be seen by the results presented in Table 5 the combined variable D H was most closely associated with the dependent variables. Tree diameter squared (D2) was the next most highly correlated of the independ-ent variables. Tree age (AGE) and height to live crown (HLC) were most poorly correlated with the tree and component dry-weights. A plotting of the dependent variables over D2H revealed with the exception of merchantable stem dry-weight (Y^), slight curvilinear relationships and increases in the variances of the dependent variables as the magnitude of the independent variable increased. In order to obtain a linear relationship and uniform variance along the regression line both the dependent and independent variables were transformed to logarithms. Table 6 and Figures 4 to 18 present the allometric equations developed by the analysis. Of the dependent variables analyzed, only five (dry stem weight (Y 3), dry merchantable stem weight (Y^), dry needle weight (Yg), dry root plus stump weight (Y 8), and dry root weight (Yg))were obtained by direct measurement. The remaining four dependent variables (dry total tree weight (Y]_) , dry above-ground weight (Y2) , dry bark weight (Y5) , and dry branch weight (Y7)) were determined indirectly and, consequently, i t is. very likely that their variability was reduced somewhat. There-fore i t is probable that the standard errors of estimate and the spread between the maximum and minimum values of these estimated variables, are low. The problem of obtaining systematic underestimates by using logarithmic regression equations has been well documented by Zar (1968), Baskerville (1970 and 1972), and Crow (1971). As can be seen by the aggregate deviations (AD) presented in Table 6 this varied from less Table 6. Dry-weight allometric relationships for 100-year-old lodgepole pine trees. Dependent Intercept Regres. Independent No. r2 Sy.x CF AD variable (lb.)1 coeff. var.(in. ft.) observ. (lb.) Average stand density (Stands 1 and 2 pooled) Logi o Yi -0.966 0.997 Logio D2H 72 0.987 32.0 1.003 -0.002 Logi o Y2 -0.998 0.985 Logi o D2H 85 0.985 28.1 1.004 -0.005 Logi o Y3 -0.889 0.938 Logio D2H 85 0.980 25.8 1.005 -0.000 Y- -12.626 0.071 D2H 85 0.979 27.4 1.000 0.000 Logi o Ys -2.584 1.140 Logi o D2H 85 0.849 11.6 1.053 -0.054 Logi o Y6 -2.951 1.148 Logi o D2H 85 0.875 4.2 1.057 -0.027 Logi o Y7 -4.126 1.509 Logi o D2H 85 0.860 11.6 1.145 -0.104 Logio Y8 -1.880 1.022 Logi o D2H 72 0.936 12.7 1.013 -0.024 Logic. Y9 -2.104 1.070 Logi o D2H 72 0.923 13.4 1.017 -0.030 High stand density (Stand 3) Logi o Yi -0.578 0.887 Logi o D2H 221 0.981 3.8 1.025 -0.038 Logi o Y2 -0.749 0.923 Logi o D2H 221 0.984 2.8 1.021 -0.021 Logi o Y3 -0.823 0.901 Logi o D2H 221 0.988 1.7 1.015 -0.013 Logi o Y6 -2.108 1.023 Logi o D2H 221 0.876 0.8 1.236 -0.151 Logi o Y7 -1.806 1.023 Logi o D2H 221 0.876 1.5 1.236 -0.150 Logio Y8 -1.123 0.806 Logi o D2H 221 0.908 1.6 1.053 -0.085 See Table 4 for definition of abbreviations. 21 F i g u r e 4. T O T A L T R E E W E I G H T - O H A L L O M E T R Y O F • A V E R A G E S T A N D D E N S I T Y 1 100-YEAR-OLD L O D G E P O L E P I N E . F i g u r e 3 . T O T A L A B O V E - G R O U N D C O M P O N E N T W E I G H T -D 2 H A L L O M E T R Y O F ' A V E R A G E S T A N D D E N S I T Y 1 1 0 0 - Y E A R - O l D L O D G E P O L E P I N E . i J J U 9 | 0 Y ) > 0 . « 9 7 I Sr,-330 lb. l i t « , 0 O 5 H » • 7 J *>> i n 3 l t . 1 - 0 . 9 6 6 6 v.* • 0.9W t o j ^ o V ( r'"0.9aj «"»S ».l lb. (9.0 %\ »3«.l-< 99« IT* (KI/11.1 0Vl (in.1-HI F i g u r e 6. S T E M W E I G H T - D H A L L O M E T R Y O F ' A V E R A G E S T A N D D E N S I T Y * 100" Y E A R -O L D L O D G E P O L E P I N E . / 1 ^ 1 0 Y J " 0»3» l o g » 3» 0 .980 «• >'•• ">•« p ^ H l i , »5 3 l l . ) - o.»«» 10 o3* F i g u r e 7. M E R C H A N T A B L E S T E M W E I G H T - D ^ H R E L A T I O N S H I P O F ' A V E R A G E S T A N D D E N S I T Y ' 1 0 0 - Y E A R - O t D L O D G E P O L E P I N E . £ too VA- 0 .071 D 3 H [ i n * l l . ) - 1 J . 6 2 6 r3" 0 . 9 7 9 n « as S,..'"' lb. a » o woo 6ooo eooo u o o o 12.000 u o c o O'H t i - 3 i . . ] 22 Figure 8. S T E M B A R K W E I G H T - D ^ H A L I O M E T R Y O F L O D G E P O L E P I N E . Figure 9. N E E D L E W E I G H T - 0 H A L I O M E T R Y O F " A V E R A G E S T A N D D E N S I T Y ' 100 - Y E A R -O L D L O D G E P O L E P I N E . 23 ngw* 12. tOOT WEIGHT - D H ALLOMETRY Of 'AVERAGE STAND DENSITY' 100-YEAR-OLD lOOGEFOLE PINE. 24 Figure 1 3 . TOTAL TREE WEIGHT-D2H ALLOMETRY OF 'HIGH STAND DENSITY' 100"YEAR-OLD LODGE POLE FINE. 2 I | I 4k 4k *4F^ 4k * A A Jr f 41 A * u>QioVft* 7 Log, < -pi ,02M[iii.a-#t.J-Q 578 $ r . - 3.8 lb. 18.3% J F i g u r e 14. T O T A L A B O V E - G R O U N D C O M P O N E N T W E I G H T -D 2 H A L L O M E T R Y O F " H I G H S T A N D D E N S I T Y ' W O - Y E A R - O L D L O D G E P O L E P I N E . 3EK, «r 1 o i J < A 4 4k A> * 4k / . 4k I O S X J ' J " 0 " 0.749 »3'0.984 n-5 R . - 2.8 ft. ( 14.7%; 0 5 M [m. J - l l j F i g u r e 1 5 . S T E M W E I G H T - D 2 H A L L O M E T R Y O F " H I G H S T A N D D E N S I T Y ' 1 0 0 - Y E A R - O L D L O D G E -P O L E P I N E . » K i ! 4k 4k i 4k 4k 4 r 4k U e „ T , «0 •01 log ^ O'M - 0 823 4k jt .'-0.988 n "22 F i g u r e 1 6 . N E E D L E W E I G H T - D 2 H A L L O M E T R Y O F * H I G H S T A N D D E N S I T Y ' 1 0 0 - Y E A R - O L D L O 0 G E P O L E P I N E . * t Z 0'H tm?- H . J 023 to. lnDJH - 2.108 .'-0.876 S Y. „•<>.« o. [5i< i> -231 •*>] A A / 4 » . / k A "iv. •* A A A 4k 4k . A Ay A A V A * A *m A / % a * A A A AygOkA 4 A * / * A W f e A - A i / 7- 1*r • • A A \ A A A A 4b AAA A A A \ A k A n 25 f i g u r e 1 7 . B R A N C H W E I G H T - D 2 H A L L O M E T R Y O F ' H I G H S T A N D D E N S I T Y ' 1 0 0 - Y E A R - O l D I O D C E P O L E P I N E . , ' . 0 v . -Ij' 1.0 •76 1.3 lb. 23 l o s ^ D ^ M -1 „ -111 [ 5 1 . 7 % ] B06 * ft A * A ft * * * ' A, * A / A i • i F i g u r e 1 8 . S T U M P P L U S R O O T W E I G H T - D 2 H A L L O M E T R Y O F ' H I G H S T A N D D E N S I T Y ' 1 0 0 " Y E A R - O L D L O D G E P O L E P I N E . i r'- 0.901 TOOT b. [40. LOO^ D'H - I.IV I A L A . *AT * A A * A A * A 1 * «k 4ft A 0JH [ i n . 3 l l j 26 than 0.1 percent to 10.4 percent for the trees from average density stands to from 1.3 percent to 15.1 percent in the high density stands. These results suggest that for some variables (Yj, Y2, and Y 3 ) this underestimate may be ignored. In other instances, particularly branch dry-weight ( Y 7 ) , this systematic bias is considerable and cannot be overlooked. A comparison between the bias correction factors (CF) and the aggregate deviations (AD) presented in Table 6 indicates that in most cases the application of the correction factor would have resulted in an under-estimate of less than 4 percent. In the high density stand the application of the correction factor would have resulted in an over-estimate of approximately 8.5 percent for needle and branch dry-weights. This may be due to the high variability in these weights (coefficients of variation of 149 percent), and a high variability even at a given tree size (Figures 13 and 18). The total standing crop dry-weight and its distribution by components for the three stands (Table 7) was determined from the equations and correction factors listed in Table 6 and measurements of diameter at breast height, and tree height of a l l of the trees in each plot. The weights of the spruce trees present in the plot were measured and are included in Table 7. Although the number of spruce trees present was small, separate equations were derived by the same methods used for the pine trees and these equations are presented in Appendix II. 27 Table 7. Dry weight of tree components and total standing crop for three stands of 100-year-old lodgepole pine. Component STAND 1 STAND 2 STAND 3 (oven-dry pounds per acre) Needles 11,619 (4.4%) 13,324 (6.5%) 7,688 (7.6%) Branches 15,209 (5.8%) 22,589 (11.0%) 15,398 (15.1%) Stem 193,045 (73.4%) 136,134 (66.1%) 61,816 (60.8%) Total above l'-stump 222,613 (84.7%) 173,267 (84.1%) 83,207 (81.9%) Roots plus stump 39,203 (14.9%) 31,585 (15.3%) 19,228 (18.9%) Total a l l components 262,941 206,090 101.637 1 The figures are not additive because they were determined from independent least-square logarithmic regression equations. As can be seen from the results presented in the preceding table (Table 7) there is a marked difference in the distribution of dry-weight by components in the three sample stands. In Stand 2, which had been thinned previously, a greater proportion of the total dry-weight was contained in the foliage and branches than in its unthinned counterpart (Stand 1). This suggests that the thinning resulted in crown expansion at the possible expense of stem wood production. Although the mean dry weight of needles of 1.55 pounds per tree observed in Stand 3 was considerably less than that observed in Stands 1 and 2 (11.39 and 45.94 pounds per tree, respectively), the greatest propor-tion of crown materials per acre (Table 7) was supported by the highly suppressed trees in Stand 3. Because the high stand density probably resulted in a low light intensity within Stand 3, a greater proportion of foliage was probably necessary to ensure that the compensation point, at which photosynthesis equalled respiration, was attained. As a result of this intense competition for light, l i t t l e energy was avail-able for wood production. The relationships between photosynthesis 28 and l i g h t i n t e n s i t y are discussed by Kramer and Kozlowski (1960) , Nichiporovich (1967) , Wassink (1968) , and Monsi (1968). Component dry-weights of 100-year-old lodgepole pine stands Table 8 presents the stand c h a r a c t e r i s t i c s used as independent v a r i a b l e s i n the an a l y s i s of estimated t o t a l standing crop of 100-year-old lodgepole pine stands from A l b e r t a . Table 8. S t a t i s t i c a l c h a r a c t e r i s t i c s of the independent v a r i a b l e s from eighty-eight stands of 100-year-old lodgepole pine. Independent Mean Standard Minimum Maximum v a r i a b l e dev. value value Age (AGE) (yr.) 98.0 6.0 91.0 110.0 AI (AI) (%) 100.0 9.2 80.0 126.0 K l (Kl) ( f t . ) 56.8 8.2 44.0 81.0 Number of stems/acre (N) 884.1 402.1 260.0 2,140.0 Average diameter (D) (in.) 6.0 1.4 3.6 9.5 Basal area/acre (BA) (s_q.ft.) 162.9 28.2 108.6 222.8 Average stand height (H) ( f t . ) 52.2 11.3 24.1 72.9 Basal area times ave. height (BA-H) ( f t . 2 - f t . ) 8,531.4 2,433.5 3,198.1 14,347.7 Basal area divided by ave. height (BA/H) ( f t . 2 / f t . ) 3.3 0.9 1.8 5.5 Ave. diameter divided by ave. height (D/H) ( i n . / f t . ) 0.11 0.02 0.09 0.15 The estimated component dry-weights per acre used as the dependent v a r i a b l e s i n the a n a l y s i s of t o t a l standing crop of 100-year-old lodgepole pine stands from A l b e r t a are presented i n Table 9. 29 Table 9. Statistical characteristics of the dependent variables from eighty-eight stands of 100-year-old lodgepole pine. Dependent Mean Standard Minimum Maximum variable (lb./ac.) dev. value value Needle dry-weight (Yio) 6,612 1,974 3,622 11,266 Branch dry-weight (Yu) 9,154 4,189 3,719 19,894 Stem dry-weight (Y12) 135,163 30,977 82,969 214,073 Above-ground dry-weight (Y13) 152,621 36,935 93,643 246,076 Root plus stump dry-weight (YiO 27,203 6,893 16,279 44,454 Total tree dry-weight (Y15) 180,590 44,349 109,914 292,460 Weights obtained from least-square logarithmic equations and therefore are not additive. Table 10 presents the simple correlation coefficients between the dependent and independent variables of the 88 stands of 100-year-old lodgepole pine from Alberta. Table 10. Simple correlation coefficients (r) between several stand characteristics and stand component dry-weights for eighty-eight stands of 100-year-old lodgepole pine. Independent Dependent variables variables1 Yio Y11 Y12 Y13 Ym Yis AGE 0.262 0.314 0.175 0.201 0.218 0.207 AI 0.407 0.238 0.556 0.521 0.494 0.513 Kl 0.595 0.656 0.444 0.492 0.523 0.502 N -0.463 -0.643 -0.170 -0.255 -0.313 -0.275 D 0.762 0.872 0.537 0.606 0.652 0.622 BA 0.601 0.372 0.796 0.751 0.716 0.740 H 0.754 0.756 0.642 0.683 0.707 0.691 BA-H 0.949 0.816 0.976 0.981 0.979 0.981 BA/H -0.277 -0.366 -0.117 -0.164 -0.197 -0.175 D/H 0.053 0.257 -0.176 -0.116 -0.072 -0.101 p.05 signif. level r i > 8 6= 0.212 1 See Table 8 for definition of abbreviations. 2 See Table 9 for definition of abbreviations. 30 As can be seen from the results presented in Table 10, component dry-weights per acre, with the exception of branch dry-weight per acre (Yii), are most closely associated with the combined variable of stand basal area times the average height of the trees in the stand (BA'H). Similar high correlations were observed between stand volume and the combined variable (BA«H) over a wide range of site and age conditions from approximately 3,200 lodgepole pine sample plots in British Columbia (Smith, 1972) and from 820 lodgepole pine sample plots in Alberta (Johnstone, 1972). Branch dry-weight per acre was most closely associated with mean stand diameter. The relationships between biomass per acre and the combined variable are presented, for each component, in Figures 19 to 24. As expected the dry weights of needles and branches decreased with increasing numbers of stems per acre (Table 10). It was somewhat surprising however to note that this decrease in component dry-weight per acre with increasing number of stems held true for a l l components. This is probably due to the fact that although number of stems and basal area were positively correlated (r = 0.345), number of stems was negatively correlated with mean stand diameter and mean stand height (r = -0.848 and -0.687, respectively). The negative influence of number of stems on mean stand height was sufficient to result in a negative correlation (r = -0.315) between number of stems and the combined variable (BA'H). It is undoubtedly for these same reasons that the crowding ratios, BA/H and D/H, were poorly correlated with a l l of the dependent variables. In addition, number of stems (N) was negatively correlated (r = -0.663) with conventional site index (KI) and as such the reduction of component biomass with increasing number of stems may also be a reflection of a reduction in site quality as well as an increase in competition. Figure 25 presents the relationship between total standing crop and number of stems. The 31 H o w 19. THf RELATIONSHIP IETWEEN ESTIMATED DRY NEEDLE WEIGHT Ft• ACM AND STAND 1ASAL AREA TIMES MEAN STAND HEIGHT FO* STANDS OF KX>-TEA«-OtO IODGEFOIE FINE. IMOO titm 20. THE RELATIONSHIP BETWEEN ESTIMATED DRY IRANCH WEIGHT FER ACRE AND STAND IASAL AREA TIMES MEAN STAND HEIGHT FOR STANDS OF 100-YEAR-OLD IODGEFOIE FINE. t 1 I i %» H (h.'-ti.) a.ooo 13.000 • A A (It'-tt. ) r^ura at. THE RELATIONSHIP MTWEEN ESTIMATED DRY STEM WEIGHT PER ACRE AMD STAND BASAL AREA TIMES MEAN STAND HEIGHT FOR STANDS Of 100 - YEAR - OLD LODGEPOLE PINE. vn- it.mi + n.43 •* ft .'•MM m-tt tow upoo Figur»22. THE RELATIONSHIP BETWEEN ESTIMATED DRY ABOVE "GROUND WEIGHT PER ACRE AND STAND BASAL AREA TIMES MEAN STAND HEIGHT r*OR STANDS OF KW-YEAR-OLD LODGEPOLE PINE. 130.000 r ) 3 - l i . j j o » tutt i* fi •A A (ti.7H.) 32 33 Figure 25. THE RELATIONSHIP BETWEEN ESTIMATED DRY TOTAL TREE WEIGHT PER ACRE AND NUMBER OF STEMS PER ACRE FOR STANDS OF 100"YEAR-OLD LODGEPOLE PINE. 3 0 0 . 0 0 0 -2 6 0 . 0 0 0 . i 2 2 0 . 0 0 0 . I I A A A A O 1 8 0 . 0 0 0 A A * 1 1 4 0 . 0 0 0 -A A A A A A A A A 1 0 0 . 0 0 0 . 2 0 0 6 0 0 ljOOO N u m b e r o f S t e m s 1 . 4 0 0 1 . 8 0 0 2 . 2 0 0 34 heterogeneity of variance displayed in this relationship (Figure 25) was characteristic of the relationships between the dry weight per acre for each component and number of stems. Multiple regression equations relating the influences of site productivity, number of stems per acre, and basal area per acre to total tree biomass per acre are presented in Table 11. The regression statistics presented are: regression coefficients (B_^ ) and their signi-ficance (F .), number of observations (No.).coefficients of determinations xi (R2), and standard errors of estimate (Sy.x). The results presented in Table 11 further demonstrate the inverse relationship between number of stems and total standing crop. These results also indicate that on a given site a stand of a small number of large trees will contain a greater total standing crop than a stand of a large number of small trees when the basal area of the two stands is equal. These relationships were characteristic for a l l components. . Table 11. Regression statistics for evaluating the influences of site quality, number of stems per acre and basal area per acre on the total standing crop of 100-year-old lodgepole pine stands. Dependent Independent variables and their significance No. R Sy.x variable (lb./ac. (Xi = N, X2 = BA, X3 = AI or Kl) (lb.) ) Regression coefficients and F-ratios 80 81 FX! 82 Fx2 83 Fx3 X3 = AI Yis -120,237 -68.9 520.8 1,308.6 811.0 1,485 .1 119.4 88 0.945 10,596 Y15 -3,337 -66.4 203.1 1,489.0 503.7 88 0.867 16,393 Y15 -107,221 1,000.5 72.9 1,247 .8 11.9 88 0.603 28,266 Y 1 5 -68,396 -43.6 21.7 2,875 .4 48.8 88 0.413 34,383 X3 = Kl Y 1 5 -146,179 -34.5 100.6 1,416.1 1,461.8 2,228 .0 194.3 88 0.960 9,060 Yis -216,015 1,286.3 632.6 3,292 .4 349.4 88 0.911 13,351 Yis -3,337 -66.4 203.1 1,489.0 503.7 88 0.867 16,393 Y15 -5,165 11.5 0.7 3,091 .6 21.0 88 0.259 38,635 p. 01 s ignificance level Fj_ 8-= 6.95 ^1 t 8 5 = • 6.95 36 Conclusions It is possible to make reliable estimates of the dry weights of the various components of mature lodgepole pine trees from the relatively simple measurements of tree height and diameter at breast height. It is probable that these estimates would be improved had the quantity of bark and branch components been obtained by direct measure-ment. Systematic underestimates will result i f the estimates are based on logarithmic equations (Table 6). This result supports the findings of Zar (1968), Baskerville (1970, 1972) and Crow (1971). The magnitude of the underestimate varies with components and is greatest for the crown components. Meyer's (1944) method for overcoming this problem is generally satisfactory but may, as in the case of the crown components, result in a slight overestimate (Table 6). With the exception of the crown components (i.e., needles and branches) the relative magnitude of the errors are small and for pratical purposes can be ignored. These results suggest therefore, that reliable estimates of component or total tree dry-weight per unit area can be obtained using double sampling techniques. It has long been recognized that the height growth of some tree species is influenced by extremes in stand density. This is part-icularly true of lodgepole pine trees (Smithers, 1961; Alexander, Tackle and Dahms, 1967). In this study similarities in climatic and edaphic factors suggest that the productivity of the tree sample stands (Stands 1, 2, and 3) should be similar. Assuming this to be true, as can be seen from the mean height data presented in Table 2, any attempt to assess site productivity in terms of conventional height-age relationships may 37 r e s u l t i n erroneous conclusions. However, from the r e s u l t s presented i n Table 7, comparisons of p r o d u c t i v i t y i n terms of t o t a l tree biomass are also unadvisable. C o r r e c t i o n f o r the obvious omission of the biomass contributed by l e s s e r ground vegetation would not improve the estimate s i g n i f i c a n t l y because i n a l l three stands t h i s vegetation was n e g l i g i b l e . These r e s u l t s suggest therefore, that one must be cautious i n comparing stand dry-matter production when extremes i n stand density are encountered or when a major stand disturbance has occurred a f t e r the age at which the stand can f u l l y compensate f o r that disturbance. Although the biomass estimates of the 88 stands of 100-year-old lodgepole pine represent, at best, a f i r s t approximation (Table 9), the a n a l y s i s does suggest that, with the possible exception of branch biomass per acre, r e l i a b l e estimates of component biomass can be obtained using the combined v a r i a b l e of basal area per acre times mean stand height (Table 10). The strong inverse c o r r e l a t i o n between branch biomass and numbers of stems per acre and the strong postive c o r r e l a t i o n of branch biomass and average stand diameter r e f l e c t s the r e l a t i v e i n t o l e r a n t nature of the species. The r e s u l t s i n d i c a t e that the amount of f o l i a g e i s c l o s e l y associated with dry-matter production (Table 10) thus sup-po r t i n g the r e s u l t s of Mar:Moller (1947), B a s k e r v i l l e (1965a), and S t i e l l (1966). Unlike MarrMoller (1947) and S t i e l l (1966), a constant f o l i a g e biomass was not apparent, although the range i n density examined i n t h i s study was r e l a t i v e l y narrow. S i m i l a r l y , no maximum t o t a l standing crop was observed. Mar:Moller (1947) hypothesized that a f t e r f u l l s i t e occupancy had been achieved growth becomes s t a t i c and changes i n stand density only a l t e r the d i s t r i b u t i o n and not the amount of growth. The r e s u l t s presented i n Table 11 tend to refute t h i s hypothesis f o r lodgepole pine. The r e s u l t s presented therein demonstrate that, on a given s i t e 38 and at a given stand basal area, component and total tree biomass per acre decreases as the number of stems increases. This undoubtedly reflects the fact that as the numbers of stems increase, live crown length decreases and thus the amount of foliage per acre decreases, resulting in a lower total growth. This further demonstrates the need to account for both stocking (area occupancy) and stand density (crowding within the area stocked) when considering the growth and yield of lodgepole pine stands. 39 CHAPTER I I I DRY-MATTER COMPARISONS BETWEEN YOUNG LODGEPOLE PINE AND POPULUS STANDS Introduction As demonstrated i n the preceding chapter of t h i s thesis considerable a t t e n t i o n has been devoted to studies of the t o t a l p r o d u c t i v i t y , i n terms of dry-matter production, of many f o r e s t eco-systems. S u r p r i s i n g l y , r e l a t i v e l y few studies (Ovington, 1957; Tadaki et a l . , 1962; B a s k e r v i l l e , 1965a, 1966; Ando, 1965; K i r a and Shidei, 1967; B e l l a and J a r v i s , 1967; Satoo, 1967; Osborn, 1968; Peterson et a l . , 1970; Johnstone, 1971; Olson, 1971; Steinbeck and May, 1971; Moir and Fra n c i s , 1972; P o l l a r d , 1972; Hegyi, 1972) have been devoted to comparing the dry-matter production of a s i n g l e species over a range of s i t e and/or stand conditions. Fewer yet are the number of studies which compare the dry-matter production of d i f f e r e n t species grown on s i m i l a r s i t e s (Mar:Moller, 1947; Ovington, 1956; Assmann, 1961; Post, 1970). The objec t i v e of t h i s study was to compare, f o r two d i f f e r e n t areas, the above-ground t o t a l standing crop of a f u l l y -stocked stand of young lodgepole pine with a f u l l y - s t o c k e d stand of Populus species of a s i m i l a r age and grown under s i m i l a r edaphic and c l i m a t i c f a c t o r s . 40 Methods and M a t e r i a l s The data f o r t h i s study were gathered from four, one hundred square metre (approximately f o u r t i e t h - a c r e ) sample p l o t s located i n two areas near Fox Creek, A l b e r t a i n the Boreal b i o c l i m a t i c zone (Rowe, 1959). In Area 1 (North Latitude 54°14', West Longitude 116°17*), paired lodgepole pine (Plot 4) and trembling aspen (Populus tremuloides Michx.) (Plot 5) p l o t s were est a b l i s h e d on an upper slope of an e l e v a t i o n of about 2,200 f e e t . The s o i l i s an o r t h i c grey wooded o v e r t i l l , and the dominant understory vegetation was Viburnum-Linneae. A small number of white spruce and willow ( S a l i x L.) trees were also present. A l l pine and aspen trees were about 23 years o l d . In Area 2 (North Latitude 54°17', West Longitude 116°10'), paired lodgepole pine (Plot 6) and balsam poplar (Populus balsamifera L) (Plot 7) p l o t s were established on a lower slope at an e l e v a t i o n of about 2,265 f e e t . The s o i l i s a gleyed grey wooded over l a c u s t r i n e c l a y , and the dominant understory vegetation was Lonicera-Heracleum. A small number of white spruce, willow and white b i r c h (Betula p a p y r i f e r a Marsh.) trees were also present. The pine and poplar trees were about 25 years o l d . The c h a r a c t e r i s t i c s of the stands, based on p l o t measurements, are presented i n Table 12. 41 Table 12. Characteristics of the four lodgepole pine and Populus stands. Characteristic Plot 4 Plot 5 Plot 6 Plot 7 (l.pine) (t.aspen) (l.pine) (b.poplar) Living stems per acre 6,515 7,972 3,683 2,671 Basal area per acre (sq.ft.) 135.7 121.3 142.0 169.9 Mean dbh (in.) 1.9 1.5 2.5 3.0 Range of dbh (in.) 0.4-3.8 0.3-3.8 0.7-4.8 0.2-6.0 Mean height (ft.) 19.1 20.2 28.6 28.8 In the pine plots (Plots 4 and 6) the diameter and height of each tree was measured. The age at stump height was determined for approximately twenty pine trees per plot. The pine trees were cut at 1.0 foot above ground and the fresh weights of the stems, branches greater than 2 cm. in diameter, branches less than 2 cm. in diameter plus foliage and attached dead branches were determined. The above-stump fresh weights of a l l living minor species and a l l dead standing trees were obtained by species. In addition, ten pine trees (covering the range of tree sizes encountered in each plot) were felled in each area, from which the follow-ing measurements were determined: a. Diameter at breast height outside bark (D) b. Diameter at crown base outside bark (DCB) c. Total height (H) d. Crown length (CL) e. Crown width (CW) f. Age (AGE) g. Stump fresh-weight h. Stem fresh-weight i . Fresh, and dry weights of branches greater than 2 cm. 42 j . Fresh, and dry weights of branches less than 2 cm. plus foliage k. Dry weight of foliage 1. Dry weight of branches less than 2 cm. m. Fresh, and dry weights of attached dead branches n. Fresh, and dry weights of the bark and wood of discs collected at ground level, stump height, breast height, and at 6.0-foot intervals above ground level in the stem, and at random from six living branches greater than 2 cm. Pine component fresh- and dry-weights, and fresh- and dry-weight allometric relationships were obtained using methods similar to those outlined in Chapter 2 of this thesis. In addition to the measured plot fresh-weights, estimated plot fresh- and dry-weights were obtained from tree height and diameter data and the following regression models: Y = Bo + 3iD + g2D2 (for dry-weights) Y = Bo + BiD2H (for fresh-weights) Localized plot dry-weights were obtained by multiplying the estimated plot dry-weight by the ratio of observed fresh-weight to estimated fresh-weight for each component. In the Populus plots (Plots 5 and 7) data were gathered follow-ing the aggregate harvest method, for immature stands between the ages of 10 and 40 years, outlined by Peterson (1970). Measured total standing crop fresh-weights were provided by Peterson (1972). Estimated plot fresh- and dry-weights were calculated from plot stand tables and regression equations derived for young aspen trees in Manitoba and Saskatchewan (Bella, 1968). For this purpose aspen and poplar component weights were estimated from the same equations based on the following multiple regression model: 43 Y = Bo + 3lD2 + B2D2H Localized plot dry-weights were obtained by multiplying the estimated plot dry-weight by the ratio of observed fresh-weight to estimated fresh-weight for each component. Estimated dry-weights for minor species other than pine or Populus species were obtained by multiplying the measured fresh-weight by the dry-weight to fresh-weight ratio of pine for coniferous minor species or the dry-weight to fresh-weight ratio of aspen or poplar for deciduous minor species. Analysis The regression techniques used in this analysis are the same as those outlined in the preceding chapter of this thesis. Tree component fresh- and dry-weights, in pounds, were used as dependent variables with several tree characteristics (Table 13) used as independent variables. The following were used as dependent variables in the regression analyses of the tree and component weights (lb.): a) Dry stump weight (Yi): the dry weight of that portion of the stem from ground level to 1.0 foot above ground level. b) Dry bole weight (Y2) : the dry weight of the stem between a 1.0-foot stump and the top of the tree. c) Dry stem weight (Y3): the dry weight of the stem between ground level and the top of the tree (Yi + Y 2). d) Dry branch weight (YO: the dry weight of a l l living branches. e) Dry needle weight ( Y 5 ) : the dry weight of a l l green needles. f) Dry dead branch weight (Ye): the dry weight of a l l dead 44 branches. g) Above-ground living dry-weight ( Y 7 ) : the dry weight of al l living components above ground level (Y3 + Yii + Y 5 ) . h) Fresh stem weight (Ys) : the fresh weight of the stem between ground level and the top of the tree. i) Fresh crown weight (Y9): the fresh weight of a l l living branches plus green needles. j) Above-ground living fresh-weight (Yio): the fresh weight of a l l living components above ground level (Ys + Y9 ) . Results and Discussion Table 13 presents the means, standard deviations, and minimum and maximum values of the independent variables used in the regression analyses. 45 Table 13. Statistical characteristics of the independent variables from twenty young lodgepole pine trees. Independent variable Mean Standard deviation Minimum value Maximum value Diameter (D) (in.) 2.1 0.8 1.0 3.4 Height (H) (ft.) 25.4 5.2 17.5 33.6 Crown length (CL) (ft.) 11.1 3.5 5.5 16.9 Crown width (CW) (ft.) 2.5 1.1 1.0 5.4 Age (AGE) (yrs.) 25.6 2.8 19.0 29.0 Diameter at crown base (DCB) (in.) 1.6 0.7 0.6 3.0 Height to live crown (Ht.LC) (ft.) 14.3 2.8 9.5 18.6 Diameter squared (D ) (in.2) 4.9 3.3 1.0 11.6 Diameter squared times height (D2H) (in.2ft .) 139.1 112.6 17.5 388.4 Dry-weight relationships The means, standard deviations, and maximum and minimum values of the dry-weight dependent variables are presented in Table 14. Table 14. Statistical characteristics of the dry-weight dependent variables from twenty young lodgepole pine trees. Dependent variable Mean Stand. Min. Max. dev. value value Dry stump wt. (Yi) (lb.) 0.9 0.6 0.2 2.0 Dry bole wt. (Y2) (lb.) 10.7 7.9 1.4 29.8 Dry stem wt. (Y3) (lb.) 11.7 8.4 1.6 31.9 Dry branch wt. (YO (lb.) 0.9 0.8 0.1 3.0 Dry needle wt. (Ys) (lb.) 1.2 1.1 0.1 4.0 Dry dead branch wt. (Ye) (lb.) 0.5 0.3 0.1 0.9 Above-ground living dry-wt. (Y7) (lb.) 13.7 10.2 1.8 38.9 46 Simple correlation coefficients (r) between the independent and dependent variables are shown in Table 15. Of the independent variables analyzed, tree diameter squared (D2) was most closely assoc-iated with four of the dependent variables (above-ground living dry-weight, stem dry-weight, bole dry-weight, and needle dry-weight); tree diameter (D), tree diameter squared times height (D2H) and crown length (CL) were most closely associated with stump dry-weight, branch dry-weight, and dead branch dry-weight, respectively. Age and height-to-live-crown were most poorly correlated with the component weights. Table 15. Simple correlation coefficients (r) between component dry-weights and several tree characteristics of twenty young lodgepole pine trees. Tree Component dry-weights  characteristics1 Yj Y2 Y3 Y,, Y5 Y6 Y7 D 0.975 0.961 0.964 0.915 0.930 0.829 0.961 H 0.791 0.854 0.852 0.831 0.772 0.836 0.846 CL 0.839 0.875 0.874 0.846 0.794 0.841 0.868 CW 0.856 0.857 0.859 0.826 0.865 0.767 0.861 AGE 0.627 0.562 0.568 0.494 0.542 0.575 0.562 DCB 0.975 0.974 0.976 0.940 0.948 0.835 0.975 Ht.LC 0.424 0.499 0.495 0.491 0.445 0.507 0.492 D2 0.973 0.983 0.985 0.959 0.960 0.781 0.985 D2H 0.935 0.980 0.980 0.975 0.948 0.762 0.980 . 05 signif. level r1 18= 0.444 See Tables 13 and 14 for description of abbreviations. Because of the high correlations among the independent variables, l i t t l e improvement in prediction reliability could be achieved using multiple regression equations. In order to estimate the dry weights of needles, branches, 47 stems, and l i v i n g tree above-ground i n the sample p l o t s the following regression model was se l e c t e d : Y = 30+ SiD + 3aD 2 Although the independent v a r i a b l e tree diameter (D) only contributed s i g n i f i c a n t l y to the equation f o r dry branch weight (Table 16) i t was retained i n the equations f o r needle dry-weight, stem dry-weight, and above-ground l i v i n g dry-weight to ensure the a d d i t i v i t y of the equations. Table 16 and Figures 26 to 29 show the regression equations used to estimate the lodgepole pine component dry-weights of the sample p l o t s . Table 16. Regression equations, and regression c o e f f i c i e n t s i g n i f i c a n c e t e s t s (Fx.), derived from twenty young lodgepole pine trees used to estimate sample p l o t l i v i n g component dry-weights. Dependent Regression c o e f f i c i e n t s and F - r a t i o s R 2 Sy.x (lb.) v a r i a b l e ( l b . ) 1 Bo Bi Fxi Fx 2 Y 3 1.074 -1.922 0.56 2.962 24.37 0.971 1.51 Yu 0.669 -0.999 8.81 0.464 35.04 0.947 0.20 Y S 0.322 -0.669 1.73 0.460 15.04 0.928 0.30 Y 6 2.065 -3.590 1.36 3.886 29.40 0.972 1.80 p.05 s i g n i f i c a n c e l e v e l Fj 17= 4.45 p.01 s i g n i f i c a n c e l e v e l F : l7= 8.40 1 See Table 14 f o r d e s c r i p t i o n of abbreviations. Fresh-weight r e l a t i o n s h i p s The means, standard deviations, and minimum and maximum values of the fresh-weight dependent v a r i a b l e s are presented i n Table 17. 48 F i g u r e 2 6 . T H E R E L A T I O N S H I P B E T W E E N S T E M D R Y -W E I G H T A N D T R E E D I A M E T E R O F Y O U N G L O D G E P O L E P I N E T R E E S . F i g u r e 2 7 . T H E R E L A T I O N S H I P B E T W E E N B R A N C H D R Y -W E I G H T A N D T R E E D I A M E T E R O F Y O U N G L O D G E P O L E P I N E T R E E S . S,.," V5I Ib.d2.99b) 0.971 n • 20 Y4-0.449 - 0.999D+0.464 0 ir,- 0.20 0.(22.2%) l*-0.947 n-20 F i g u r e 2 8 . T H E R E L A T I O N S H I P B E T W E E N N E E D L E D R Y -W E I G H T A N D T R E E D I A M E T E R O F Y O U N G L O D G E P O L E P I N E T R E E S . 7-6-5-O (in.) F i g u r e 2 9 . T H E R E L A T I O N S H I P B E T W E E N A B O V E -G R O U N D D R Y - W E I G H T A N D T R E E D I A M E T E R O F Y O U N G L O D G E P O L E P I N E T R E E S . 70 60 50 0 1 , , , , , 0 1 2 3 4 5 6 D (in.) 49 Table 17. S t a t i s t i c a l c h a r a c t e r i s t i c s of the fresh-weight dependent v a r i a b l e s from twenty young lodgepole pine trees. Dependent Mean Stand Min. Max. v a r i a b l e dev. value value Fresh stem weight (Ya) (lb.) 24.7 19.3 3.3 70.0 Fresh crown weight (Y 9) (lb.) 4.0 3.6 0.3 13.9 Above-ground l i v i n g fresh-weight 28.7 22.8 3.6 83.9 (Yio) (lb.) Simple c o r r e l a t i o n c o e f f i c i e n t s between the independent and dependent v a r i a b l e s are presented i n Table 18. Table 18. Simple c o r r e l a t i o n c o e f f i c i e n t s (r) between component fresh-weights and several tree c h a r a c t e r i s t i c s of twenty young lodgepole pine trees. Tree c h a r a c t e r i s t i c s 1 Component fresh--weights1 Y 8 Y 9 Yio D 0.958 0.931 0.956 H 0.877 0.810 0.869 CL 0.871 0.825 0.866 CW 0.825 0.861 0.832 AGE 0.560 0.521 0.556 DCB 0.955 0.953 0.957 Ht.LC 0.545 0.478 0.536 D 2 0.988 0.968 0.988 D 2H 0.996 0.970 0.995 .05 s i g n i f . l e v e l r i , i 8 = 0.444 See Tables 13 and 17 f o r d e s c r i p t i o n of abbreviations. Unlike the r e s u l t s f o r the component dry-weights (Table 15), the r e s u l t s presented i n Table 18 i n d i c a t e d that combined v a r i a b l e D 2H i s most c l o s e l y associated with the dependent v a r i a b l e s tested. Figure 30 to 32 graph-i c a l l y present the best simple l i n e a r regression equations developed i n 50 Figure 30. THE RELATIONSHIP BETWEEN STEM FRESH-WEIGHT A N D D 2 H OF Y O U N G LODGEPOLE PINE TREES. 0 5 0 1 0 0 1 5 0 2 0 0 2 5 0 3 0 0 3 5 0 4 0 0 4 5 0 D 2 H ( i n . 2 - f t . ) Figure 31. THE RELATIONSHIP BETWEEN C R O W N FRESH-WEIGHT A N D D 2 H OF Y O U N G LODGEPOLE PINE TREES. Figure32. THE RELATIONSHIP BETWEEN A B O V E - G R O U N D FRESH-WEIGHT A N D D 2 H OF Y O U N G LODGEPOLE PINE TREES. 4 5 0 D 2 H ( i n . 2 f t . ) 51 the analyses. These equations (Figure 30 to 32) were used to estimate sample p l o t l i v i n g component fresh-weights. Above-ground organic matter production of paired lodgepole pine and Populus stands of s i m i l a r ages on s i m i l a r s i t e s Table 19 presents a comparison of the measure above-ground fresh-weights by species of the paired sample p l o t s . Table 20 presents a comparison of the estimated above-ground dry-matter contained by the l i v i n g components of trees i n the paired sample p l o t s . Table 20. A comparison of estimated above-ground dry weight (lb./ac.) of l i v i n g components i n paired lodgepole pine and Populus stands of s i m i l a r ages on s i m i l a r s i t e s . „ .. AREA 1 AREA 2 Components Plot 4 Plot 5 P l o t 6 Plot 7 ( l . p i n e ) (t.aspen) (l.pine) (b.poplar) Major Species: stem 50,202 27,588 69,368 53,368 branch 5,293 4,363 7,140 5,408 f o l i a g e 7,310 2,319 9,503 2,476 Minor Species: b. poplar ( a l l components) 667 10,953 t. aspen ( a l l components) 3,881 963 1. pine ( a l l components) 12,575 w. spruce ( a l l components) 1,555 3,775 93 willow ( a l l components) 1,307 709 2,363 w. b i r c h . ( a l l components) Major species subtotal Minor species s u b t o t a l T o t a l a l l species 62,805 7,410 70,215 34,270 28,012 62,282 86,011 6,641 92,652 61,252 61,252 52 Table 19. A comparison of measured fresh weights (lb./ac.) of paired lodgepole pine and Populus stands of s i m i l a r ages on s i m i l a r s i t e s . Components AREA 1 AREA 2 Plot 4 P l o t 5 P l o t 6 Plot 7 (l.pine) (t.aspen) (l.pine) (b.poplar) Major Species: l i v i n g : stem branch >2 cm. diam. branch <2 cm. diam. ( i n c l . f o l i a g e ) dead: branches on l i v i n g trees standing trees Minor Species: l i v i n g : b. poplar t. aspen 1. pine w. spruce willow w. b i r c h dead: 1. pine t. aspen ( a l l components) ( a l l components) ( a l l components) ( a l l components) ( a l l . components) ( a l l components) ( a l l components) ( a l l components) 105,967 58,551 152,546 111,616 218 166 24,792 13,266 32,598 14,667 4,588 1,943 1,406 8,296 3,238 2,792 1,447 3,223 23,454 24,061 7,218 1,519 5,059 1,365 5,160 8,822 2,023 200 6,900 1,351 1,022 379 L i v i n g : Major species t o t a l a l l species t o t a l Dead: a l l species t o t a l 130,759 72,035 185,144 125,449 146,501 128,287 199,326 126,449 6,531 6,035 13,982 2,752 53 The results presented in Tables 19 and 20 suggest that, for lodgepole pine at least, Area 2 appears to be somewhat higher in site quality than Area 1, although the latter supported about 77 percent more trees. Reasons for the apparent reversal of this trend in the case of Populus stand are not clear but may be related to the following: a) The predominant Populus species in Area 1 was trembling aspen and the predominant Populus species in Area 2 was balsam poplar which have different productivity potentials on different sites (i.e., Area 2 was a much moister site than Area 1 and as such was unsuited for the establishment and growth of trembling aspen). b) Although fully-stocked, the Populus stand in Area 2 contained only 41 percent as many trees as did the stand in Area 1, and therefore, may not have utilized the site as fully. c) Unlike Area 2, a successional stage had been reached in Area 1 wherein the minor species present in the stand were making a large contribution to the total standing crop. In both areas the lodgepole pine outproduced the Populus species in terms of total above-ground standing crop dry-weight (1.13:1 in Area 1 and 1.51:1 in Area 2). In terms of major species distributions by components, as would be expected, a smaller proportion of total above-ground dry-weight is contained in the stems and branches of pine trees compared to Populus species (Table 21). 54 Table 21. Proportion of t o t a l above-ground dry-weight by components of major species i n study areas. Proportion (%) of t o t a l above-ground dry-weight Component AREA 1 AREA 2 Plot 4 (l.pine ) P l o t 5 (t.aspen) Plot 6 (l.pine) Plot 7 (b.poplar) Stem Branch Foliage T o t a l * 79.9 8.4 11.6 100.0 80.5 12.7 6.8 100.0 80.7 8.3 11.0 100.0 87.1 8.8 4.0 100.0 l May not be add i t i v e due to rounding e r r o r s . Conclusions This study demonstrates that the p r o d u c t i v i t i e s of d i f f e r e n t species are not constant among themselves and that considerable f l u c t u a t i o n s can be expected w i t h i n species from s i t e to s i t e . These r e s u l t s support the findings of Mar:Moller (1947), Ovington (1956), Assman (1961), and Post (1970) . The a b i l i t y of coniferous stands to out-produce hardwood stands, i n terms of above-ground dry-matter production, i s demonstrated i n Table 20. This greater a b i l i t y of c o n i f e r s to produce organic matter i s probably a t t r i b u t a b l e to: a) The greater amount and b e t t e r means of d i s p l a y i n g photo-synthetic materials by c o n i f e r s (Ovington, 1956; Kramer and Kozlowski, 1960). b) The a b i l i t y of c o n i f e r s to photosynthesize f o r a longer period of time during the year (Ovington, 1956; Kramer and Kozlowski, 1960). 55 c) The greater root penetration and thus greater site utilization by conifers (Ovington, 1956). d) The more primitive water-conductive system of conifers (Assman, 1961) , and e) The generally lower nutritional requirements of conifers and pines in particular (Rennie, 1956). The comparisons made in this study are limited to one point in time. Because the Populus stands arise from suckers they may, at younger ages, out-produce pine stands, which originate from seed, as was observed in comparisons between mountain maple (Acer spicatum Lam.) and balsam f i r (Abies balsamea (L.) Mill.) by Post (1970). 56 CHAPTER IV VARIATIONS IN STEM GROWTH AS RELATED TO SEVERAL CROWN CHARACTERISTICS OF 100-YEAR-OLD LODGEPOLE PINE TREES Introduction Growth and the resultant dimensional characteristics (most notably taper and form) of trees have long been of interest to foresters. Mensurationally, interest has arisen because as Munro (1970) pointed out, the ultimate success of any forest inventory depends upon the accuracy and precision of estimation of whole and partial tree stem volumes. The desire to obtain straight, single-stemmed, low tapered trees having a minimum of knots while maintaining moderately rapid growth rates has had a marked influence on past and present genetic selection and silvicultural practice. In addition, such basic biological considerations as how trees grow and what factors control this growth have stimulated investigations in this field. Generally, in temperate climates, a tree increases in diameter annually. This increase is the result of the laying down of an unevenly distributed sheath of xylem (and, to a much lesser degree, phloem) cells, by cambial i n i t i a l s . At the risk of oversimplifying radial growth patterns the following 'rules of thumb' most often apply. In open-grown trees, with crowns extending to their bases, radial growth increased from apex to butt. In thrifty dominant trees (forest-grown) radial growth increased from the apical tip to a point within the crown in the vicinity 57 of the branch whorl supporting the most f o l i a g e . Radial growth then decreases or remains constant throughout the branchless bole, and then increases again near the butt. In suppressed forest-grown trees maximum r a d i a l growth occurs nearer the top and then progressively decreases towards the base. Under conditions of severe competition (crowding) r a d i a l growth may almost disappear near the base (missing or discontinuous r i n g s ) . In an attempt to r a t i o n a l i z e t h i s uneven d i s t r i b u t i o n of r a d i a l growth, s e v e r a l theories have been postulated. The following i s a summary of the major theories. 1. Hartig's n u t r i t i o n a l theory wherein growth was r e l a t e d to the balance between t r a n s p i r a t i o n and a s s i m i l a t i o n . The i n t e n s i t y of t r a n s p i r a t i o n and the quantity of food a v a i l a b l e f o r stem growth at any point i n the crown i s proportional to the amount of f o l i a g e above that point. 2. Schwendener's mechanistic theory i n which bole form i s dependent upon i n t e r n a l (the weight of the stem i t s e l f ) and external (wind) forces, which cause stresses i n the stem, which i n turn stimulate the cambium. Radial growth at any point i s , therefore, pro-p o r t i o n a l to the magnitude of stress at that point. 3. Jaccard's water condu c t i v i t y theory which was predicated on the b e l i e f that the conductivity of water between the roots and the crown through recent annual rings influences the c r o s s - s e c t i o n a l area of the stem. Further, dead branches reduce the conductive capacity of these rings and therefore, i n order to maintain the conductive capacity, the c r o s s - s e c t i o n a l area must increase enough to balance losses r e s u l t i n g from dying branches. 4. The hormonal theory a t t r i b u t e s the uneven d i s t r i b u t i o n of r a d i a l 58 growth to the uneven d i s t r i b u t i o n of growth substances (which o r i g i n a t e i n the elongating buds) along the bole. 5. The pipe model theory was developed by Shinozaki et a l . (1964). The amount of f o l i a g e (photosynthetic) e x i s t i n g above any l e v e l i n the stem i s proportional to the c r o s s - s e c t i o n a l area of the stem plus branches (non-photosynthetic) at that l e v e l . Consequently, changes i n the amount of photosynthetic organs above any given l e v e l i n the bole would be r e f l e c t e d by prop-o r t i o n a l changes i n the c r o s s - s e c t i o n a l area of the non-photosynthetic organs at that l e v e l . Comprehensive reviews of these theories have been presented by Onaka (1950, a and b ) , Young and Kramer (1952), Farrar (1961), Larson (1963), Shinozaki et a l . (1964), H a l l (1965), and Heger (1965). Although the p h y s i o l o g i c a l processes which c o n t r o l form are not yet f u l l y understood, i t i s apparent that the l i v e crown i s basic to a l l of these stem form theories. The purpose of t h i s study i s to in v e s t i g a t e some of the with i n and among tree v a r i a t i o n s i n the growth of lodgepole pine trees, and to r e l a t e these v a r i a t i o n s to some crown c h a r a c t e r i s t i c s . Methods and Materials Data c o l l e c t i o n The data were gathered from 20 even-aged (100-year-old) lodgepole pine trees growing i n Stands 1 and 2 described i n Chapter 2 of t h i s t h e s i s . Ten trees (Trees numbered from 1 to 10) were grown i n an undisturbed stand 59 (Stand 1) and the remaining trees (Trees 11 to 20) were grown i n a thinned stand (Stand 2). Sample trees were selected to cover a wide range of diameter c l a s s e s . Discs, about one inch t h i c k , were sawn from the stem at 1.0 foot above ground (stump h e i g h t ) , at breast height, at 9.0 feet above ground, and at 8.0-foot i n t e r v a l s thereafter to the base of the l i v e crown (lowest branch supporting l i v i n g needles). One-inch thick d i s c s were also sawn from the stem at 2.0-foot i n t e r v a l s from the top of the tree to the base of the l i v e crown. The number of di s c s c o l l e c t e d varied from tree to tree depending upon the height and the l i v e crown length of the trees. In t o t a l 358 di s c s were c o l l e c t e d . In the f i e l d , tree number and c o l l e c t i o n height were recorded on each d i s c . No d i s c measurements were made at that time. The f o l i a g e -bearing twigs from each 2.0-foot i n t e r v a l of the stem were clipped from the l a r g e r branch pa r t s , put into separate burlap sacks, and weighed. The sacks were placed i n a drying shed and needle dry-weights were obtained by means i d e n t i c a l to those o u t l i n e d i n Chapter 2 of t h i s t h e s i s . No t o t a l weight was measured f or the branch parts. In the laboratory, the diameter i n s i d e and outside bark of each a i r - d r i e d d i s c were obtained to the nearest 1/10 inch with a diameter tape. For each d i s c , four measurements (to the nearest 1/100 millimetre) were taken of the l a s t 5-year r a d i a l growth increment using an Addo-X machine at a magnification of 125X. The f i r s t measurement was taken on a l i n e of the mean radius and the remaining measurements were taken at 90-, 180-, and 270- degree angles to t h i s f i r s t measurement. The study was r e s t r i c t e d to the most r e c e n t l y completed 5-year growth period because t h i s represents the length of needle r e t e n t i o n f o r t h i s species and because changes i n the 60 crown c h a r a c t e r i s t i c s of the trees p r i o r to t h i s 5-year period would have made the v a l i d i t y of the analyses le s s c e r t a i n . Using the average of the four r a d i a l measurements, the diameter of each s e c t i o n f i v e years p r i o r was determined. The d i f f e r e n c e between the c r o s s - s e c t i o n a l area of each d i s c determined from the present diameter and the diameter 5 years before represented the 5-year growth i n c r o s s - s e c t i o n a l area at each point i n the stem. Section volume growth w i t h i n each tree was determined by mul t i p l y i n g the average of the c r o s s - s e c t i o n a l growth of two consecutive discs times the distance between these d i s c s . Tree volume growth equals the sum of the volume growth from a l l of the sections w i t h i n each tree. The t o t a l age of each d i s c was also determined. In a d d i t i o n , a complete stem a n a l y s i s ( i n c l u d i n g earlywood and latewood measurements) was c a r r i e d out f o r a l l rings of each d i s c c o l l e c t e d from one tree grown i n each of the two stands. Tree 9 from Stand 1 and Tree 11 from Stand 2 were selected because the diameters at breast height of these two trees corresponded most c l o s e l y with the mean diameters at breast height of the ten trees sampled from each respective stand. The data from each of the two trees were c o l l a t e d and summarized from each tree using a s p e c i a l computer program f o r tree r i n g a n a l y s i s . Analysis Two s t a t i s t i c a l techniques were used to analyze wit h i n and among tree v a r i a t i o n s i n stem growth. The f i r s t technique, an analysis of variance, was used to te s t d i f f e r e n c e s between p l o t s , between trees within p l o t s , between sections (these d i f f e r e n c e s were furth e r broken down into d i f f e r e n c e s between crown-formed wood and c l e a r bole-formed wood and into 61 differences between sections within the bole and within the crown). Finally, for each plot, differences between sections were broken down into differences between bole and crown, and into differences between sections within the bole and within the crown. In order to carry out. this analysis i t was necessary to compare section measurements taken at the same actual or the same relative positions i n each tree. This was done by using five (1.0-foot, 4.5-foot, .9.0-foot, 17.0-foot, and 25.0-foot) measurements from the clear-bole, and five (the base, one-quarter, one-half, three-quarters, and at the top of the l i v e crown) measurements from the crown. The second s t a t i s t i c a l technique, regression analysis, was used to study within and among tree patterns of radial, cross-sectional area, and section volume growth. Three separate analyses, based on the multiple regression program and procedures reported by Kozak and Smith (1965) , were used for this purpose. The f i r s t analysis incorporated a l l of the observations, whereas the second analysis was limited to observations taken solely within the crown and the third analysis was based on measure-ments from the clear bole. A fourth multiple regression analysis was used to relate tree volume growth and volume growth per unit of foliage to 2 several tree characteristics. R values were calculated as outlined on page Results and Discussion Profile diagrams which i l l u s t r a t e the radial growth at each sampling point, the foliage dry-weight in the 2.0-foot interval above each sampling point, and the cumulative (total) foliage dry-weight above each sampling point, are presented for each tree i n Appendix III. An irregular pattern of foliage distribution was noted for most trees. 62 Table 22 presents the means, standard deviations, and minimum and maximum values of the 20 sample trees. Table 22. Characteristics of the twenty, 100-year-old lodgepole pine sample trees. Characteristics Mean Standard Minimum Maximum deviation value value Tree height (H) (ft.) 60. 2 8.1 43. 5 77. 6 Dbh (D) (in.) 7. 2 2.3 3. 2 12. 9 Diam. crown base (DCB) (in.) 5. 1 2.0 2. 2 11. 0 Dry foliage weight (DFW) (lb.) 14. 8 11.3 1. 0 44. 1 5-yr. <radial growth (cm.) 0. 319 0.134 0. 038 0. 708 5-yr. 'cross-sectional area growth (cm. ) 10. 61 8.11 0. 328 63. 40 5-yr. tree volume growth (TVG) (cu.ft.) 0. 694 0.525 0. 041 2. 191 Section age (yr.) 49. 1 23.3 8. 0 101. 0 Tree volume growth/unit foliage wt. (TVG/DFW) (cu.ft./lb.) 0. 048 0.010 0. 036 0. 070 As can be seen from the preceding table, the sampling covered a wide range of tree and crown sizes. Table 23 presents the simple correlation coefficients between several tree characteristics of the 20 sample trees. 63 Table 23. Simple c o r r e l a t i o n c o e f f i c i e n t s (r) between several tree c h a r a c t e r i s t i c s of the twenty, 100-year-old lodgepole pine sample t r e e s . H D DCB DFW TVG D 2 D 2H TVG/DFW H 1.000 0.906 0.877 0.798 0.762 0.873 0.875 -0.143 D 1.000 0.966 0.939 0.924 0.980 0.959 -0.133 DCB 1.000 0.911 0.911 0.970 0.964 -0.115 DFW 1.000 0.977 0.952 0.934 -0.168 TVG 1.000 0.941 0.922 0.006 D 2 1.000 0.994 -0.132 D 2H 1.000 -0.139 TVG/DFW 1.000 . 05 s i g n i f . l e v e l r x 1 Q= 0.444 See Table 22 for d e s c r i p t i o n of abbreviations. As can be seen from the r e s u l t s presented i n the preceding table (Table 23), there i s a high degree of c o r r e l a t i o n between many of the tree character-i s t i c s . Foliage dry-weight i s most highly c o r r e l a t e d with diameter at breast height squared (D 2) and t h i s r e l a t i o n s h i p i s presented i n Figure 33. A s i m i l a r ranking of c o r r e l a t i o n c o e f f i c i e n t s between f o l i a g e dry-weight and the tree c h a r a c t e r i s t i c s studied occurred when logarithmic transform-ations were c a r r i e d out. This d i f f e r s from the r e s u l t s reported for broad-leaved species ( A t t i w i l l , 1966; K i r a and Sh i d e i , 1967; Peterson et a l . , 1970) wherein dry f o l i a g e weight was most hig h l y c o r r e l a t e d with stem diameter at the base of the l i v e crown. Tree volume growth f or the past 5-year period i s most hig h l y c o r r e l a t e d with dry f o l i a g e weight (Figure 34), which as stated previously i s hig h l y c o r r e l a t e d to tree s i z e . In an attempt to determine the r e l a t i v e e f f i c i e n c y of volume growth, the r a t i o of volume •growth to amount of f o l i a g e (TVG/DFW) was re l a t e d to several tree char-a c t e r i s t i c s . The r e s u l t s presented i n Table 23 and Figure 35 i n d i c a t e that there i s a negative but n o n - s i g n i f i c a n t r e l a t i o n s h i p between volume growth e f f i c i e n c y and tree s i z e . 64 Figure 33. THE RELATIONSHIP BETWEEN NEEDLE DRY-WEIGHT AND TREE DIAMETER FOR THE 20.100-YEAR-OLD LODGEPOLE PINE TREES. 0 2 4 6 • 10 12 14 Figure 34. THE RELATIONSHIP BETWEEN PAST 5-YEAR VOLUME GROWTH AND PRESENT NEEDLE DRY-WEIGHT OF 20. 100-YEAR-OLD LODGEPOLE PINE TREES. I.4-. 0 1 i 1 1 i r — i 0 10 20 30 40 30 AO OfW (lb.) Figure 35. THE RELATIONSHIP BETWEEN VOLUME GROWTH EFFICIENCY (cu. ft. / lb. DRY FOLIAGE) AND TREE DIAMETER FOR 20.100-YEAR-OLD LODGEPOLE PINE TREES. | OJ o ? 1 -2 4 4 8 ro ra 14 65 Table 24 presents the results of the analyses of variance of within and among tree variations in radial, cross-sectional area, and section volume growth. It is of interest to note the disparity in the A.N.O.V.A. test for differences between plots. These results indicate that although there was no significant difference between plots in terms of radial growth, significant differences were observed between plots for cross-sectional area and section volume growth. This is undoubtedly due to the fact that the sample trees chosen from Plot 1 (mean dbh of sample trees = 6.8 inches) were somewhat smaller than the sample trees selected from Plot 2 (mean sample tree dbh =7.9 inches). With the exception of the test already stated, a l l of the results of the remaining tests were in agreement. Significant differences were found between trees within plots, indicating that regardless of plot, larger trees exhibit larger growth. Significant differences were also indicated between sections. Section averages indicate that there is a decrease in radial growth from the top to the base of the tree, with a slight increase at the base. As expected, this trend is the complete reverse of the trend for the average cross-sectional area and section volume growth. These sectional variances, when further partitioned, indicate that radial growth is significantly higher in the crown than in the clear bole (the reverse being true of cross-sectional area and section volume growth), and further that there are significant differences between sections when compared at the same relative positions in the crown and clear bole. Table 24. Analyses of variance of radial, cross-sectional area, and volume growth measured at 5 positions within the crown and 5 positions within the clear-bole of ten, 100-year-old lodgepole pine trees grown in a thinned stand and ten, 100-year-old lodgepole pine trees grown in an unthinned stand. Source df Radial growth Cross-sectional area growth Section volume growth MS F MS F MS F Stands 1 568 0.18ns 174 9.33** 4,307,300 5.72* Trees within stands 18 147,900 46.36** 541 28.93** 10,113,000 13.42** Sections Crown vs. bole Sections within position (crowns vs. bole) 9 1 8 53,802 283,430 25,090 16.87** 88.85** 7.87** 350 1,714 180 18.73** 91.60** 9.62** 25,195,000 189,581.000 4,646,700 33.43** 251.55** 6.17** Stand X section (interaction) 9 Stand X crown vs. bole 1 Stand X sections within position (crowns vs. bole) 8 2,394 146 2,675 0.74ns 0.05nS 0.84nS 6 0.3 7.0 0.34ns o.oins 0.35nS 407,760 1,526,600 268,100 0.54ns 2.03ns 0.36nS Error 162 3,190 19 753,650 Total 199 ** Significant at the 0.01 probability level. * Significant at the 0.05 probability level. Not significant. ON 67 Table 25 presents some simple c o r r e l a t i o n c o e f f i c i e n t s between a l l of the s e c t i o n growth measurements i n the tree and several tree and s e c t i o n c h a r a c t e r i s t i c s . Table 25. Simple c o r r e l a t i o n c o e f f i c i e n t s (r) between 358 measurements of r a d i a l , c r o s s - s e c t i o n a l area, and s e c t i o n volume growth, and several tree and s e c t i o n c h a r a c t e r i s t i c s . C h a r a c t e r i s t i c C o r r e l a t i o n c o e f f i c i e n t s (r)  Radial growth Cross-sectional Section volume (R. Gr.) area growth growth (S.V.Gr.) (C.S.A.Gr.) Tree height (H) 0.443 0.562 0.311 Dbh (D) 0.618 0.701 0.395 Section height •(h) 0.426 -0.270 -0.525 Section diameter (d) 0.063 0.819 0.749 Section age (A) -0.343 0.417 0.583 Foliage weight i n 2 feet above (F) 0.496 0.003 -0.280 Cumulative f o l i a g e weight above (CFW) 0.347 0.884 0.678 Height from top (HT) -0.240 0.529 0.679 Percent of t o t a l height (PH) 0.321 -0.418 -0.600 Form (d/D) -0.341 0.429 0.566 p.05 s i g n i f . l e v e l r1 3 5 6 = 0.105 The preceding r e s u l t s (Table 25) i n d i c a t e that, i n general, growth, be i t r a d i a l , c r o s s - s e c t i o n a l or volume, i s most c l o s e l y associated with tree s i z e . The r e s u l t s further i n d i c a t e the following trends. Radial growth decreased from the top to the base of the tree. The reverse of t h i s trend i s true f o r c r o s s - s e c t i o n a l area and se c t i o n volume growth. These r e s u l t s were as expected and support the r e s u l t s of the analyses of variance. Further, the c o r r e l a t i o n s suggest that r a d i a l growth i s more a function of p o s i t i o n within the tree than c r o s s - s e c t i o n a l area growth, which i s most 68 c l o s e l y associated with the s i z e of the tree at the points of growth. I t i s of i n t e r e s t to note that r a d i a l growth i s more c l o s e l y associated with the amount of f o l i a g e i n the 2.0-foot i n t e r v a l immed-i a t e l y above the point of growth than i t i s with the t o t a l amount of f o l i a g e above the point of growth. The opposite of t h i s a s s o c i a t i o n i s true of c r o s s - s e c t i o n a l area and s e c t i o n volume growth. This d i f f e r e n c e undoubtedly r e f l e c t s the fact that the cumulative f o l i a g e weight i s probably as much a function of tree s i z e as i t i s of p o s i t i o n w i t h i n the tree. From the preceding a n a l y s i s i t i s obvious that i n order to s u c c e s s f u l l y p r e d i c t e i t h e r growth pattern one must f i r s t account f o r v a r i a t i o n s due to tree s i z e . The following three equations appear to be best f o r r e l a t i n g r a d i a l , c r o s s - s e c t i o n a l area and s e c t i o n volume growth i n e n t i r e stems to the quantity of f o l i a g e . Logio R.Gr. = 2.095+0.0346 Logio F + 0.054 D R 2 = 0.46 n = 358 Logio C.S.A.Gr. = 0.114 + 0.598 Logio CFW + 0.033 D R 2 - 0.86 n = 358 Logio SVGr. = 2.122 + 0.183 L o g 1 0 CFW + 0.076 D R 2 = 0.523 n = 358 The f i r s t equation accounted f o r only 46 percent of the v a r i a t i o n i n r a d i a l growth, the second equation accounted f o r 86 percent of the v a r i a t i o n i n c r o s s - s e c t i o n a l area growth, and the t h i r d equation accounted f o r 52 per-cent of the v a r i a t i o n i n s e c t i o n volume growth. Table 26 presents some simple c o r r e l a t i o n c o e f f i c i e n t s between measurements of the r a d i a l , c r o s s - s e c t i o n a l area, and s e c t i o n volume growth of crown-formed wood and c l e a r bole-formed wood, and sev e r a l tree and 69 crown c h a r a c t e r i s t i c s . Table 26. Simple c o r r e l a t i o n c o e f f i c i e n t s (r) between 245 measurements of r a d i a l , c r o s s - s e c t i o n a l area, and se c t i o n volume growth i n crown-formed wood and between 113 measurements of r a d i a l , c r o s s - s e c t i o n a l area, and sect i o n volume growth i n bole-formed wood, and several tree and s e c t i o n c h a r a c t e r i s t i c s . C o r r e l a t i o n c o e f f i c i e n t s (r) C h a r a c t e r i s t i c Crown-formed wood Bole--formed wood R. Gr. C.S.A.Gr . S.V. Gr. R.Gr. C.S.A. Gr. S.-V.Gi Tree height (H) 0.454 0.639 0.620 0. 406 0.604 0.589 Dbh (D) 0.623 0.764 0.744 0. 621 0.792 0.753 Section height (h) 0.481 -0.075 • -0.126 0.027 • -0.150 0.024 Section diameter (d) 0.150 0.872 0.892 0. 585 0.835 0.669 Section age (A) -0.310 0.451 0.494 0. 108 0.289 0.096 Foliage weight i n 2 f t . above (F) 0.495 0.247 0.218 — — — Cumulative f o l i a g e weight above (CFW) 0.290 0.910 0.912 0. 714 0.866 0.820 Height from top (HT) -0.117 0.670 0.706 0. 223 0.494 0.338 Percent of t o t a l height (PH) 0.228 -0.527 • -0.570 -0. 040 -0.236 -0.070 Form (d/D) -0.263 0.480 0.541 0. 092 0.271 0.044 p.05 s i g n i f . l e v e l r l j X 1 1 = 0.187 v1 2 i * 3 = 0.126 The r e s u l t s presented i n Table 26 generally agree with those presented i n Table 25, suggesting that there i s l i t t l e advantage i n d i f f e r e n t i a t i n g between bole-formed and crown-formed wood when studying r a d i a l cross-s e c t i o n a l , or s e c t i o n volume area growth. The r e s u l t s i n Table 26 tend to support the trends suggested e a r l i e r i n t h i s study wherein r a d i a l growth appears to be a function of p o s i t i o n within the tree whereas cross-s e c t i o n a l area and sec t i o n volume growth i s not only a function of p o s i t i o n but also of the s i z e of the tree at the point where the growth i s measured. The following equations appear to be the most s a t i s f a c t o r y f o r r e l a t i n g the r a d i a l and c r o s s - s e c t i o n a l area growth of crown-formed and 70 clear bole-formed wood to the amount of foliage. Crown-formed wood: Logio R.Gr. = 2.229 + 0.145 Logio F + 0.038 D R2 = 0.46 n = 245 Logio C.S.A.Gr. = 0.101 + 0.532 Logio CFW + 0.040 D R2 = 0.87 n = 245 Log10 S.V.Gr. = 1.897 + 0.130 Logio CFW + 0.086 D R2 = 0.75 n = 245 Clear bole-formed wood: Logio R.Gr. = 1.909 + 0.573 Logio CFW - 0.326 HT R2 = 0.71 n = 113 Logio C.S.A.Gr. = 0.147 + 0.890 Logio CFW - 0.005 h R2 = 0.89 n = 113 Logio S.V.Gr. = 2.285 + 1.038 Logio CFW - 0.024 d R2 = 0.74 n = 113 For the two trees (Trees 9 and 11) subjected to complete stem analysis, earlywood formed during the last five years generally increased to a maximum occurring near the top of both trees. In Tree 9 this maximum was observed at the highest sampling point (two feet from the top) and in Tree 11 the maximum occurred six feet from the top at the base of the section containing the greatest amount of foliage in the tree. Latewood formed during the last five years, although much more uniform than early-wood, appeared to be greatest both at the base and top of the trees. Minimum latewood formation occurred at a point eight to ten feet from the top of the tree and immediately above a point of high latewood formation. For both trees latewood to earlywood ratios decreased from the base to the top of the tree with a slight increase occurring near the top. For both trees irregularities in foliage distribution were not reflected in 71 i n e i t h e r earlywood or latewood formation. Conclusions The r e s u l t s of the analyses i n d i c a t e that r a d i a l , cross-s e c t i o n a l area, and sec t i o n volume growth are highly v a r i a b l e both within and among 100-year-old lodgepole pine trees. Thinning does not appear to have aff e c t e d the current pattern of r a d i a l growth (Table 24). This i s probably due to the l a t e age at which the stand was thinned. Kramer and Kozlowski (1960) suggested that i n older stands, wherein the r e s i d u a l trees are small crowned, increases i n photosynthesis are accompanied by increases i n r e s p i r a t i o n and the net gain i s n e g l i g i b l e . In add i t i o n , any shade needles present i n the crowns of the r e s i d u a l trees were probably injured when exposed to f u l l s u n l i g h t following thinning. Radial growth appears to decrease from the apex towards the bottom of the tree with a s l i g h t increase at the base corresponding to the butt swell. Onaka (1950a) a t t r i b u t e d t h i s phenomenon, at l e a s t i n part, to the d i s t r i b u t i o n of hormonal and food substances, which are produced high i n the crown, diminish i n quantity as they move down the crown, and accumulate at the base. Because the trees examined i n the present study cannot be considered suppressed, the r e s u l t s obtained are not i n complete agreement with the general patterns of r a d i a l growth reported i n the l i t e r a t u r e (Onaka, 1950a; Farrar, 1961; Larson, 1963; Shinozaki et_ al_., 1964) which suggest that r a d i a l growth increases from the top to approximately the point of maximum f o l i a g e , whereafter i t remains constant or decreases to the base. The low c o r r e l a t i o n s between f o l i a g e weight and r a d i a l growth within the crown (Table 26) tend to support H a l l ' s (1965) and Heger's (1965) conclusions that some factors i n a d d i t i o n to n u t r i t i o n a l gradients determine the 72 pattern of stem wood increment. However, unl i k e H a l l ' s (1965) r e s u l t s , the d i s t r i b u t i o n of growth was not c l o s e l y r e l a t e d to distance from apex although t h i s discrepancy may be due to the advanced age of the trees when the present study was c a r r i e d out. The two complete stem analyses c a r r i e d out i n the present study i n d i c a t e d d i s t r i b u t i o n s of earlywood and latewood s i m i l a r to those reported by Heger (1965). The r e s u l t s presented i n Figure 35, which suggest that the amount of t i s s u e produced per pound of f o l i a g e i s unrelated to tree s i z e (and thus crown s i z e ) , agree with the r e s u l t s presented for red pine (Pinus  resinosa A i t . ) by H a l l (1965) but disagree with those presented for black spruce (Picea mariana ( M i l l . ) BSP.) by Weetman and Harland (1964) and for balsam f i r (Abies balsamea (L.) M i l l . ) by B a s k e r v i l l e (1965a). » Because balsam f i r and black spruce are r e l a t i v e l y more shade tole r a n t than lodge-pole and red pine, small crowned trees of the former species probably have a higher proportion of shade needles which are photosynthetically more e f f i c i e n t at low l i g h t i n t e n s i t i e s (Kramer and Kozlowski, 1960). Although i n s u f f i c i e n t l y tested, the r e s u l t s presented i n Figure 34 suggest that r e l i a b l e p r o j e c t i o n s of needle dry-weights of stands and trees are obtain-able from estimates of p e r i o d i c annual volume increment. Although some s i g n i f i c a n t c o r r e l a t i o n s were found between f o l i a g e weight and growth, the implications of these r e s u l t s are unclear.. For example, although c r o s s - s e c t i o n a l area growth was h i g h l y c o r r e l a t e d with cumulative f o l i a g e weight, t h i s may merely be a r e f l e c t i o n of tree (stem) s i z e . Therefore, the r e s u l t s presented herein, by themselves, can-not be used to defend or refute any of the form theories presented i n the i n t r o d u c t i o n to t h i s chapter. Further, these r e s u l t s are of l i t t l e value i n d e f i n i n g thinning or pruning p r e s c r i p t i o n s . 73 Recognizing these limitations in the present study, i t is possible to suggest how a study, having similar objectives, should be undertaken. The first recommendation would be to limit the number of measurements taken in any one tree and to increase the number of trees studied. The results of the present study would have been much more revealing had a wider range of stand densities (from open to closed) been considered. Measurements could have been limited to five relative positions within the crown and five relative positions within the bole. This would have permitted easier analysis and a larger number of trees to have been measured for an equal amount of work. Further, had younger (less than 50 years old) , more vigorous trees been studied no doubt the results would have been clearer, more dramatic, and of greater use in planning the management of stands on the rotations likely to be used in Alberta. Such results would be of interest in studying the concepts of crown and growth distribution relationships put forward by Shea and Armson (1972). Also, recently developed X-ray methods (Parker and Henoch, 1971) should be used to determine the influence of various crown development factors on the distribution of wood substances, because differences between the specific gravities of earlywood and latewood are highly significant. 74 CHAPTER V COMPLETE-TREE UTILIZATION OF AVERAGE STAND DENSITY 100-YEAR-OLD LODGEPOLE PINE Introduction T r a d i t i o n a l l y , i n North America, the sole source of f i b r e for pulping has been the bark-free 'merchantable' bole. H i s t o r i c a l l y , research, such as that reported by Crossley (1938) and S p r o u l l e_t al_. (1957) , on the p o t e n t i a l of u t i l i z i n g other components for pulp f i b r e has been rather l i m i t e d . However, i f the projected world demand for wood-fibre products w i t h i n the next 30 years i s c o r r e c t , a shortage of as much as 100 m i l l i o n cubic metres (or 44 m i l l i o n oven-dry tons, based on an average bulk density of 25 l b . / f t . 3 ) of wood has been forecast by Keays (1970) and Keays and Hatton (1971a). In a n t i c i p a t i o n of t h i s shortage increased a t t e n t i o n has r e c e n t l y been devoted to c l o s e r and more complete u t i l i z a t i o n of f o r e s t trees and stands. To date the most comprehensive studies of complete-tree u t i l i z a -t i o n have been c a r r i e d out by Professor Harold Young and h i s colleagues at the U n i v e r s i t y of Maine (Chase et a l . , 1971; Dyer, 1967; Dyer et a l . , 1968; Young, 1964, 1965a, 1965b, 1966, 1967, 1968; Young and Chase, 1965; Young and Guinn, 1966; Young et a l . , 1964; Young et a l . , 1965). The f e a s i b i l i t y of complete-tree u t i l i z a t i o n of Canadian wood species i s being studied by the Canadian Forestry Service at i t s Western Forest Products Laboratory (Keays, 1971a,b; Keays and Hatton, 1971a,b; Hatton and Keays, 1971; Hatton and Samkova, 1972). The r e s u l t s of these studies suggest that those tree components previously considered n o n - u t i l i z a b l e and l e f t as residue 75 following logging can be converted into s a t i s f a c t o r y pulp. Table 27 presents a general summary of the pulp y i e l d and strength q u a l i t y of the various previously u n u t i l i z e d tree components. Table 27. Comparison of r e l a t i v e values of k r a f t pulp from c o n i f e r tree components expressed i n terms of the bole. Pulp properties Non-merch. Branches Root + stump top (>1 in.) (>1 in.) (>1 in.) Y i e l d at 20 permanganate no. 99% ± 1 70% 99% ± 1 Burst f a c t o r 100% ± 5 55% ± 5 85% ± 5 Tear f a c t o r 85% ± 5 75% ± 5 92.5% ± 2.5 Breaking length 100% ± 5 55% ± 5 85% ± 5 Beating time f a s t e r f a s t e r slower Source: Keays (1971a) Based on these estimates (Table 27) i t i s apparent that, i n terms of pulp y i e l d and q u a l i t y , non-merchantable tops can and should be u t i l i z e d . Small amounts of branch wood may be u t i l i z e d i f economic means can be developed to harvest, debark, and chip branches. Although the r e s u l t s i n Table 27 suggest that the root-stump component converts into s a t i s f a c t o r y pulp, u t i l i z a t i o n i s doubtful because of problems i n harvesting, cleaning, tra n s p o r t i n g , debarking and chipping t h i s component. According to Keays (1971a), the c r i t i c a l questions which r e l a t e to complete-tree u t i l i z a t i o n are: 1. How much of each component i s a v a i l a b l e f o r u t i l i z a t i o n ? 2. What q u a l i t y of pulp or other product could be obtained from each tree component? 3. How would the various tree components be extracted, trans-ported and processed? 76 4. What would be the e f f e c t of complete-tree u t i l i z a t i o n on fo r e s t growth and regeneration, on s i l v i c u l t u r a l p r a c t i c e s , on the t o t a l ecology? The purpose of the present study i s to answer, at l e a s t i n part, the f i r s t two of these questions f o r lodgepole pine, a species of great importance to the fo r e s t economies' of Al b e r t a and the I n t e r i o r of B r i t i s h Columbia. L i t t l e information pertinent to the l a t t e r two questions i s presently a v a i l a b l e , and consequently only a l i m i t e d d i s c u s s i o n of these topics i s presented. Methods and Materials Data c o l l e c t i o n Ten sample trees from Stands 1 and 2 (previously described i n Chapter 2) were used to study the pulp y i e l d and q u a l i t y of 100-year-old, forest-grown lodgepole pine t r e e s 1 . No attempt was made at randomization i n the s e l e c t i o n process, and the choice of sample trees was based on the tree's s i z e (two trees from each of the 4-, 6-, 8-, 10-, and 12-inch dbh classes were chosen) and apparent freeness from i n j u r y ( i . e . , some of the tree boles i n Stand 2 were scarred during the e a r l i e r thinning of the stand). The trees were cut at a 1.0-foot stump, and weighed with branches and f o l i a g e i n t a c t ( t o t a l above-ground fresh-weight) and with the branches and f o l i a g e removed ( t o t a l stem fresh-weight). The stem was subdivided i n t o the merchantable bole (equal to or greater than 4.0 inches i n diameter (ob)), The complete-tree u t i l i z a t i o n p o t e n t i a l of seven white spruce trees grown i n Stand 2 were reported by Keays and Hatton (1971b). 77 non-merchantable top (that p o r t i o n of the stem le s s than 4.0 inches but greater than or equal to 1.0 inch i n diameter (ob)) , and t i p (that p o r t i o n . of the stem le s s than 1.0 inch i n diameter (ob)) , and the weights of these three components were measured. The merchantable bole and non-merchantable top were each further subdivided i n t o four sections of equal length, and one-inch t h i c k d i s c s , f o r moisture content determinations (as previously outlined i n Chapter 2) were sawn from the end of each section. Unbarked wood samples for the pulp evaluation studies were obtained from each merchantable bole and non-merchantable top s e c t i o n as f o l l o w s : 1. For those sections weighing l e s s than or approximately equal to 15.0 pounds (fresh-weight) the e n t i r e s e c t i o n was taken, and 2. For those sections weighing more than 15.0 pounds ( f r e s h -weight) a sample, weighing approximately 15.0 pounds and sawn at an angle of 40° from a plane perpendicular to the tree length, was removed from the centre of each section. A more d e t a i l e d d e s c r i p t i o n of the pulp sample c o l l e c t i o n procedure i s given by Keays (1968). The branches and needles were subdivided i n t o branches 1.0 inch i n diameter and greater, and into branches l e s s than 1.0 inch i n diameter plus needles, and weighed. Discs , f o r moisture content determination, were c o l l e c t e d from the branches. A l l of the branches greater than or equal to 1.0 inch i n diameter were retained because i n a l l cases the weight of these branches was l e s s than the 60.0 pounds per tree desired for the pulp evaluation study. A l l of the needle-bearing twigs were clipped from the branches le s s than 1.0 inch i n diameter (which were then discarded), put i n t o burlap sacks and dri e d under conditions s i m i l a r to those noted i n Chapter 2. 78 The root-stump component of each tree was uprooted and washed free of s o i l . After the surface had dried the component was weighed, a l l roots less than 1.0 inch in diameter were removed and the weight of the root-stump system greater than or equal to 1.0 inch in diameter was obtained. Discs were cut from the root-stump component so that dry-weight conversions could be made. For those trees with root-stump components greater than or equal to 1.0 inch in diameter, weighing less than or equal to 60.0 pounds the entire component was retained for the pulp evaluation study. Approximately 60.0 pounds of unbarked wood samples were retained from those trees with root-stump components greater than or equal to 1.0 inch in diameter which exceeded 60.0 pounds. The pulp evaluation study samples were placed in crates and shipped to the Western Forest Products Laboratory. Some of the charac-teristics of the ten trees used in the pulp study are presented in Table 28. At the laboratory the samples were stored at a temperature below freezing. Each sample was barked by hand and slabbed on a band saw at a 40° angle to their grain to give discs 1/2 - 3/4 inch thick. The discs were reduced to chips by a laboratory chip splitter or by hand. The chips were air-dried to a moisture content of about 8 percent, and screened (using a C.J. Wennbergs chip slot screen (type KJL, standard C7)) , with only those chips 2-4 mm thick accepted for pulping. Knots, bark, and other impurities were removed from the accept chips. It was decided to limit the pulp evaluation study to the following three components: f u l l bole (merchantable bole plus non-merchantable top),branches 1.0 inch in diameter and greater, and the root plus stump 1.0 inch in diameter and greater. In order to ensure that Table 28. Characteristics of the selected 100-year-old lodgepole pine trees. Tree Age DBH Height Merch. Non-merch. No. (yr.) (in.) (ft.) bole bole (4"-lM) Component dry-weights (lb.) bark included length length Root-St. Root-St. Bole Top Top Branch Branch Dead Needles (ft.) (ft.) < 1" > 1" > 4" 4, ,-l" •<1" > 1" < 1" Branches 1 95 7.1 64.2 42.4 18.7 3.6 50.5 192.7 20.5 1.0 0.0 15.1 3.0 8.5 2 95 12.0 74.3 63.8 8.0 42.6 255.6 823.8 8.6 0.5 30.9 121.9 37.0 60.6 3 98 4.5 55.0 14.0 37.4 2.7 15.8 49.3 61.7 1.0 0.0 9.0 1.0 8.6 4 97 5.8 53.2 30.4 18.4 4.5 30.3 125.2 33.3 0.5 0.0 7.5 1.0 7.4 5 99 5.6 52.3 30.7 18.9 4.3 27.5 113.5 30.6 0.5 0.0 8.7 2.0 8.9 6 96 9.9 66.7 53.7 10.3 28.1 142.1 493.1 18.2 0.4 4.7 67.7 2.0 38.3 7 96 10.1 72.0 55.8 13.2 13.3 142.7 470.6 16.1 0.4 4.4 48.5 8.0 33.2 8 98 8.0 58.7 46.2 8.8 11.3 70.4 286.8 11.1 0.4 3.8 36.2 1.0 23.8 10 92 12.0 81.2 67.9 9.8 20.9 204.4 810.0 8.6 0.4 17.2 82.2 23.0 28.1 11 97 4.3 47.1 9.6 35.0 0.6 15.5 30.5 55.8 0.6 0.0 3.2 1.0 2.6 Mean 96. 3 79.3 62.5 41.5 17.9 13.2 95.5 339.6 26.5 0.6 6.1 40.0 7.9 22.0 St.dev. 2. 0 2.9 11.0 20.0 10.5 13.6 85.7 297.7 19.0 0.2 10.2 39.9 12.3 18.4 VO 80 the chips were representative of the f u l l bole of each t r e e , prorated amounts of chips (based on section volume w i t h i n each t r e e ) , from each sample s e c t i o n , were thoroughly blended together. P r i o r to cooking dup-l i c a t e chip moisture content determinations, using about 60 grams of a i r -dry chips per determination, were c a r r i e d out for each t r e e . The laboratory pulping equipment used i n t h i s study was a s t a i n l e s s s t e e l Weverk research d i g e s t e r with a 1.0-cubic-foot c a p a c i t y . The material to be pulped was sealed i n s t a i n l e s s s t e e l bombs (each of 735-ml capacity) which were placed i n the research d i g e s t e r . A d e t a i l e d d e s c r i p t i o n of the digester assembly was presented by Keays and Bagley (1970) . The experimental design of t h i s study required four bombs (two f o r screenings, permanganate number and y i e l d determinations, and two f o r PFI m i l l and strength determinations) from each component of each t r e e . Only f i v e of the ten sample trees contained branches greater than 1.0 inch i n diameter. In a d d i t i o n , i n order to evaluate the v a r i a t i o n i n pulp q u a l i t y w i t h i n the t r e e , two bombs (one f o r screenings, permanganate number and y i e l d , and one for PFI m i l l and stength) from each of the eight bole sample sections were cooked f o r one t r e e . Each bomb contained about 60 grams of a i r - d r i e d c h i p s . Because i t i s d e s i r a b l e to compare pulps of a uniform permanganate number (Keays et a l . , 1969; Hatton and Keays, 1970,) an exploratory cook, using chips from the branches, root-stump system and prorated bole of one tree and the pulping conditions presented i n Table 29, was c a r r i e d out to s e l e c t the e f f e c t i v e a l k a l i needed to produce an unbleached pulp with a permanganate number of approximately 20 from each component. The permanganate numbers of the pulp produced using these conditions are also presented i n Table 29. 81 Table 29. K r a f t cooking conditions used for the Exploratory cooks. F u l l bole Root-stump Branches E f f e c t i v e a l k a l i s (%) 16. ,0,17.0,18. 0 16.0,17.0,18.0 19.0,20.0,21.0 S u l f i d i t y (%) 25.4 25.4 25.4 Time to max.temp, (min.) 135 135 135 Time at max.temp, (min.) 75 75 75 Max. temp. (°C) 170 170 170 Liquor-to-wood r a t i o 4.5:1 4.5:1 4.5:1 Unbleached pulp permanganate numbers obtained 18 .7,17.2,15. 3 17.7,15.6,13.9 16.6,15.0,13.5 Based on the r e s u l t s of the exploratory cook i t was poss i b l e to estimate the e f f e c t i v e a l k a l i required to^produce an unbleached pulp with a permanganate number of approximately 20 (Figure 36). The following cooking conditions (Table 30) were used f o r a l l the subsequent cooks i n t h i s study. Table 30. Kraft cooking c o n d i t i o n used i n study to obtain unbleached pulps with permanganate numbers of approximately 20 from the three components of 100-year-old, forest-grown lodgepole pine trees. E f f e c t i v e a l k a l i : f u l l bole (%) 15.3 root-stump (%) 14.8 branches (%) 16.8 S u l f i d i t y (%) 25.4 Time to max. temp. (min.) 135 Time at max. temp. (min.) 75 Max. temp. (°C) 170 Liquor-to-wood r a t i o 4.5 Upon completion of the cooking, black l i q u o r was c o l l e c t e d from each bomb and retained f o r r e s i d u a l a l k a l i a n a l y s i s . The cooked chips of each bomb were d i s i n t e g r a t e d f o r f i v e minutes and the r e s u l t i n g pulp was thoroughly washed. The pulp to be used f o r y i e l d determination was oven-82 Figure36. THE RELATIONSHIP BETWEEN U N B L E A C H E D PULP P E R M A N G A N A T E NUMBER A N D EFFECTIVE ALKALI FOR THREE C O M P O N E N T S O F 100-YEAR-OLD L O D G E P O L E PINE. 21 r 20 -19 18 JO E 17 16 IS 14 13 j l . 14 IS 16 17 18 Effective Alkali ! % J 19 20 21 83 drie d at 105°C for a minimum of 16 hours and had reached a' constant dry-weight. The pulp to be used f o r PFI m i l l and strength determinations was a i r - d r i e d f o r a s i m i l a r time period. The pulp from each bomb was r e -d i s i n t e g r a t e d f o r f i v e minutes and the amount of r e j e c t s was determined by weighing the oven-dry material retained on a 10-cut plate of a V a l l e y laboratory f l a t screen. The screened pulp from each bomb was centrifuged f o r f i v e minutes, thoroughly mixed and conditioned to a uniform moisture content of about 8 percent. Unbleached permanganate numbers were determined on the screened pulp used f o r the y i e l d determinations following Tappi Standard T214ts-50 (using 40 ml of 0.1N KMnOu). Each determination was done i n d u p l i c a t e . Both the pH and r e s i d u a l a l k a l i (following methods s i m i l a r to those out-l i n e d by Macdonald (1969)) were determined f o r the black l i q u o r from each bomb. The screened pulp from the two bombs for each component from each tree to be used f o r the stength determinations were thoroughly com-bined. This mixture was d i v i d e d i n t o two equal a l i q u o t s , from each of which a sample of 24.0 grams O.D. was withdrawn. In some instances, p a r t i c u l a r l y i n the case of branches, a smaller sample was necessitated by low y i e l d s . Each sample was d i s i n t e g r a t e d at a consistency of about 1.2.percent u n t i l free from f i b r e bundles and c l o t s . The sample was then adjusted to a consistency of 10 percent and placed i n the PFI m i l l . The PFI m i l l methods used were s i m i l a r to those ou t l i n e d by Standard Testing Procedure PB-6 of the Pulp and Paper Research I n s t i t u t e of Canada (Pulp and Paper Research I n s t i t u t e of Canada, 1962). Two conditioning runs were c a r r i e d out before the beating of each component. The o b j e c t i v e of t h i s study was to test the strength of the unbleached pulp from the d i f f e r e n t components at a freeness of 300 ml CSF. 84 Because previous tests of lodgepole pine pulp (Keays, 1972) indicated that the strength curves are relatively flat at this freeness, i t was decided that when a freeness of 300 ml ±10 ml CSF was obtained only one PFI mill run would be carried out. If the freeness obtained was outside these limits a second PFI mill run, using the second pulp sample from the same component of the same tree, was carried out to obtain a freeness on the opposite side of 300 ml CSF from the f i r s t . In this event pulp strength properties at 300 ml CSF were obtained by interpolation. Pulp freeness was tested following Tappi Standard T227 os-58, and six hand sheets for each PFI mill run were prepared according to Tappi Standard T205 n-58. The five best hand sheets for each run were tested for breaking length, bulk, and burst and tearing strength, following Tappi Standard T220 m-60. Analysis The data were analyzed using regression (with the computer program described by Kozak and Smith (1965)), analysis of variance, and graphical techniques. Where logarithmic equations were developed the standard errors of estimate (s ) and coefficients of determination y-x (R2 or r2) were determined by the methods described previously in this thesis (Chapter 2). Unlike the results presented in the biomass section of this thesis (Chapter 2), the results presented in this section are limited to that portion of the components considered to be potentially utilizable for pulping. It is necessary therefore to redefine the components as follows: 85 1. Merchantable bole: that portion of the stem from a 1.0-foot stump to a A.0-inch top diameter (ob). 2. Non-merchantable top: that p o r t i o n of the stem from a 4.0-inch diameter (ob) to a 1.0-inch diameter (ob). 3. F u l l bole: the merchantable bole plus the non-merchantable top ( i . e . , that portion of the stem from a 1.0-foot stump to a 1.0-inch top diameter (ob). 4. Branches: those branches greater than or equal to 1.0 inch i n diameter (ob). 5. Root-stump system: the stump ( i . e . , that p o r t i o n of the stem from ground l e v e l to 1.0 foot above ground l e v e l ) plus the roots ( i . e . , that portion of the tree below ground l e v e l ) greater than 1.0 inch i n diameter (ob). In the following s e c t i o n of t h i s chapter, references to the components are to those defined above unless otherwise s p e c i f i e d . In order to eliminate e r r o r i n the comparison of y i e l d of pulps varying i n permanganate number (Hatton and Keays, 1970) an adjusted pulp y i e l d at permanganate number 20 was ca l c u l a t e d f o r a l l y i e l d data using the following formula: Y 2 0 = Y• + 3 (20.0-K ) obs. obs. where: Y 2 u = adjusted y i e l d to permanganate number 20 Y , = observed y i e l d obs. J ^obs = ° bs e r v e c* permanganate number 3 = component slope constant for the r e l a t i o n s h i p between pulp y i e l d and permanganate number (Figure 37) and equals 0.38, 0.36, and 0.43 for the f u l l b o l e , root-stump, and branch components, r e s p e c t i v e l y . The 0.38 slope observed for the f u l l bole agrees with that reported for lodgepole pine Permanganate Number 87 by Hatton and Keays (1970) . Results and Discussion The quantity of each component p o t e n t i a l l y a v a i l a b l e from the complete-tree u t i l i z a t i o n of 100-year-old lodgepole pine trees The equations presented i n Table 6 and Figures 6, 10, and 11 are of use i n obtaining gross estimates of the wood resource p o t e n t i a l l y a v a i l a b l e through complete-tree u t i l i z a t i o n . However, the estimates thus obtained are overestimates because they include such n o n - u t i l i z a b l e com-ponents as bark and portions of the components les s than 1.0 inch i n diameter. Unfortunately, s u f f i c i e n t data necessary to develop complemen-tary bark-free, oven-dry weight equations for the components, as rede-fine d on page 85 of t h i s t h e s i s , were not a v a i l a b l e . Bark-free oven-dry wood weights for each component are shown i n Table 31 expressed as percentages of the f u l l b ole. Table 31. Bark-free, oven-dry weights of the 100-year-old lodgepole pine tree components as a percentage of f u l l b oles. Tree Dbh Tree components (weight i n lb.) no. i (in.) Merch. Non-merch. F u l l Branches Root-•Stump bole bole bole Wt. % Wt. % Wt. % Wt. % Wt. % 1 7.1 175.5 90.8 17.7 9.2 193.2 100.0 0.0 0.0 43.9 22.7 2 12.0 762.8 99.1 7.1 0.9 769.9 100.0 24.5 3.2 231.1 30.0 3 4.5 45.3 45.1 55.2 54.9 100.5 100.0 0.0 0.0 14.6 14.5 4 5.8 115.2 79.4 29.8 20.6 145.0 100.0 0.0 0.0 27.3 18.8 5 5.6 103.1 79.6 26.4 20.4 129.5 100.0 0.0 0.0 24.7 19.1 6 9.9 455.6 96.6 15.8 3.4 471.4 100.0 3.7 0.8 129.6 27.5 7 10.1 436.2 96.9 13.8 3.1 450.0 100.0 3.6 0.8 127.3 28.3 8 8.0 261.6 96.5 9.4 3.5 271.0 100.0 2.9 1.1 63.4 23.4 10 12.0 754.1 99.0 7.4 1.0 761.5 100.0 12.8 1.7 187.4 24.6 11 4.3 27.9 35.7 50.3 64.3 78.2 100.0 0.0 0.0 13.4 17.7 Mean 313.7 81.9 23.3 18.1 337.0 100.0 4.8 0.8 86.3 22.6 Stan . dev. 276.9 23.2 17.3 23.2 263.8 - 8.0 1.0 78.0 5.1 88 Figure 38 i l l u s t r a t e s the r e l a t i o n s h i p between component bark-f r e e , oven-dry weight expressed as a percentage of the f u l l bole weight and tree diameter. In s e l e c t i n g an equation s u i t a b l e for describing the r e l a t i o n s h i p s for the merchantable bole and the non-merchantable top, two conditions were sought, namely: a) The equation for the non-merchantable top should equal 100 percent at a dbh (ob) of 4.0 inches (the merchantability l i m i t ) and asympototic to 0.0 percent at large diameters, and b) The equation f o r the merchantable bole should equal 0.0 percent at a dbh (ob) of 4.0 inches and asymptotic to 100.0 percent at large diameters. The equations presented i n Figure 38 s a t i s f y the above conditions. Although s i m i l a r t h e o r e t i c a l conditions undoubtedly apply to the root-stump and branch components greater than 1.0 inch i n diameter ( i . e . , the percentages f o r these components increase from 0.0 f o r small trees to a probable maxima for large trees) the range of tree s i z e s samples was too narrow to include these co n d i t i o n s . Consequently, for the root-stump and branch components, conditioning was not attempted and the equations presented i n Figure 38 were the best equations ( i n terms of standard errors of estimate and c o e f f i c i e n t s of determination) obtained. As can be seen from the r e s u l t s presented i n Table 31 and Figure 38 the quantity of wood f i b r e contained i n the non-merchantable top, branches, and root-stump system i s considerable. Because present harvesting p r a c t i c e s can r e a d i l y accommodate the e x t r a c t i o n of the non-merchantable top, t h i s component i s of immediate i n t e r e s t . The r e s u l t s presented i n Table 32 confirm those reported by Smithers (1961) and Lee (1967), and demonstrate t h a t , even i n older aged stands over a wide range of s i t e q u a l i t i e s , the Figure 38. THE RELATIONSHIP BETWEEN TREE SIZE AND OVEN-DRY, BARK-FREE COMPONENT WEIGHT EXPRESSED AS A PERCENTAGE OF THE OVEN-DRX BARK-FREE FULL BOLE WEIGHT FOR 100-YEAR-OLD LODGEPOLE PINE TREES. l o o | ( ) N M T [ % ] « 4 . 2 5 5 - 3 . 8 8 4 l o g 0 8 H - 0 . 0 1 6 6 8 t o g [ D B H - 3.99°3 OBH [in.] 90 numbers of lodgepole pine trees per acre present in the 4- to 8-inch diameter classes are substantial. It is apparent from the results presented in Figure 38 that the percentage of the full bole contained in the non-merchantable top increases very rapidly with decreasing diameter through this diameter range. It follows therefore, that the yield of wood fibre per acre can be greatly increased by harvesting to a 1.0-inch top diameter. Because the taper and form of trees are affected by stand density (Smithers, 1961; Lee, 1967) the results presented herein may only be representative of trees growing under similar stand density conditions. Table 32. Average number of stems per acre in diameter classes less than 9.0 inches dbhob from eighty-eight stands of 100-year-old lodgepole pine. Dbhob Class (in.) Total Total 1 2 3 4 5 6 7 8 <9.0 in. 1.0 in, Ave. : 20 34 100 172 171 148 107 66 819 884 Range: 0-365 0-230 0-530 0-740 0-490 15-320 10-304 0-184 110-2140 260-2140 Relative to the non-merchantable top and the root-stump system, the quantity of branches greater than 1.0 inch in diameter is small (Figure 38). According to Smithers (1961), the occurrence of large branches on lodgepole pine trees is mainly limited to trees grown in the open or in low-density stands (100 to 600 mature trees per acre). Relatively l i t t l e is known about the rooting habit of lodgepole pine trees. Horton (1958) reported changes in root development at different ages and on different soil types. Because there is l i t t l e doubt that relationships exist between rooting habit and site productivity, the results presented in Figure 38 are probably applicable only to trees growing under similar 91 site and stand conditions. Further analysis of the component weights of the individual trees examined in Chapter 2 of this thesis has been carried out by Keays (1971 b,c,d,e,f). The quality of pulp obtained from the complete-tree utilization of 100-year-old lodgepole pine trees Variation among trees The following is a discussion of the variations observed in the yield and quality of kraft pulp produced from several components of 100-year-old lodgepole pine trees. For the f u l l bole component further comparisons are made with the results reported for lodgepole pine by Hatton and Keays (1970) and Keays (1972) which were obtained under similar laboratory conditions (i.e., the same equipment, test procedures and personnel), from merchantable trees of the same age to those used in this thesis. Table 33 presents unadjusted and adjusted (permanganate number 20) pulp yield data obtained for several components for 100-year-old lodgepole pine cooked by the kraft process. 92 Table 33. Component pulp y i e l d data from k r a f t pulping for ten, 100-year-old lodgepole pine trees. Unadjusted Adjusted Component: unscreened Permanganate unscreened Screenings y i e l d (%) number y i e l d (%) % F u l l bole: Mean 47.11 20.31 46.99 0.84 Range 45.54-48.86 18.8-23.3 45.33-48.78 0.11-2.20 95% C.I. (%-width) 0.45 0.59 0.52 0.31 Reference standard 1 44.3 18.5 44.8 Root-stump: Mean 46.00 19.82 46.06 0.64 Range 41.26-47.46 18.60-21.10 41.76-47.50 0.16-1.78 95% C.I. (%-width) 0.76 0.33 0.72 0.21 Branches: 2 Mean 38.85 20.49 38.64 0.05 Range 37.68-39.93 19.60-21.80 37.50-39.84 0.00-0.20 95% C.I. (J^-width) 0.58 0.54 0.65 0.05 1 Hatton and Keays (1970) and Keays (1972). 2 Branches a v a i l a b l e from 5 trees only. The average adjusted pulp y i e l d of 46.99 percent at 20 permanganate number fo r the f u l l bole component i s somewhat higher than the adjusted pulp y i e l d of 44.8 obtained for lodgepole pine trees of a s i m i l a r age by Hatton and Keays (1970). Values of burst f a c t o r , breaking length, tear f a c t o r , and bulk at 300 ml CSF of pulp handsheets prepared from the components of 100-year-old lodgepole pine trees are summarized i n Table 34. 93 Component: Table 34. Component pulp q u a l i t y at 300 ml CSF from k r a f t pulping for ten, 100-year-old lodgepole pine trees. Unbleached pulp q u a l i t y Burst f a c t o r Breaking length (m) Tear f a c t o r Bulk (cc/gm) F u l l bole: Mean 97.2 Range 93.0-104.0 95% C.I. (%-width) 2.9 Reference standard 1 106±1 Root-stump: Mean 74.3 Range 67.1-83.2 95% C.I. (Js-width) 3.4 Branches: Mean 53.2 Range 51.6-55.0 95% C.I. (%-width) 1.6 12,648 12,150-13,480 358 14,400±350 9,952 9,080-10,990 485 7,002 6,726-7,450 361 115 104-125 5.3 105±4 120 103-143 9.0 103 100-107 3.4 1.41 1.38-1.44 0.02 1.41+0.02 1.38 1.31-1.45 0.03 1.28 1.25-1.29 0.02 1 Keays (1972). Branches a v a i l a b l e from 5 trees only. Because of dif f e r e n c e s i n beating techniques ( i . e . , a PFI m i l l was used i n t h i s study and a Valle y beater was used by Keays (1972)) the pulp q u a l i t y r e s u l t s presented i n Table 34 are not d i r e c t l y comparable. However, the s i m i l a r i t i e s between the r e s u l t s obtained and the r e s u l t s reported by Keays (1972) are s u f f i c i e n t l y close to suggest that the r e s u l t s obtained are probably representative of the species. The lower y i e l d and strength of branchwood pulp compared to f u l l bole pulp (Tables 33 and 34), as Keays (1971d) pointed out, i s probably due to a lower c e l l u l o s e content, and higher l i g n i n , mannan, pentosan and ash content of the branchwood. These d i f f e r e n c e s are a manifestation of the 94 higher percentage of compression wood i n the branches. The greater s i m i l a r i t y i n pulp y i e l d and q u a l i t y between the root-stump and f u l l bole components than between the branches and the f u l l bole i s probably a r e l e c t i o n of the greater s i m i l a r i t i e s i n s p e c i f i c g r a v i t y , f i b r e length and width, and c e l l w a l l thickness between root-stumpwood and bolewood (Keays, 1971f). Table 35 presents the simple c o r r e l a t i o n c o e f f i c i e n t s (r) between tree diameter (D) and adjusted pulp y i e l d and pulp q u a l i t y for several components of 100-year-old lodgepole pine cooked by the k r a f t process. Table 35. Simple c o r r e l a t i o n c o e f f i c i e n t s (r) between tree s i z e (D) and component unscreened pulp y i e l d (at permanganate number 20) and component unbleached pulp q u a l i t y (at 300 ml CSF) from k r a f t pulping ten, 100-year-old lodgepole pine t r e e s . Component Independent Dependent v a r i a b l e v a r i a b l e Adjusted y i e l d1 Burst factor Breaking l e n g t h2 Tear f a c t o r2 Bulk* F u l l bole: D 0.822 -0.279 -0.547 0.245 0.384 D 2 0.815 -0.332 -0.559 0.261 0.396 D 3 0.795 -0.369 -0.563 0.262 0.391 Root-stump: D -0.006 0.199 0.092 -0.005 0.050 D 2 0.057 0.201 0.095 0.015 0.110 D 3 0.108 0.203 0.098 0.040 0.164 Branches:3 D -0.368 0.089 0.500 -0.823 -0.709 D 2 -0.338 0.091 0.507 -0.804 -0.713 D 3 -0.310 0.093 0.511 -0.783 -0.714 Two observations per tree (p.05 s i g n i f . l e v e l r i > 1 8 = 0.444) One observation per tree (p.05 s i g n i f . l e v e l r i ^ s3 0.623) Branches a v a i l a b l e from 5 trees only (p.05 s i g n i f . l e v e l rj. } 3= 0.878) 95 As can be seen from the r e s u l t s presented i n Table 35 only the r e l a t i o n s h i p between the adjusted pulp y i e l d of the f u l l bole component and tree s i z e was s i g n i f i c a n t . The r e l a t i o n s h i p between the adjusted pulp y i e l d of the f u l l bole component and tree s i z e appears to be of a l i n e a r form (Table 35) and i s presented i n Figure 39. Although there i s a p o s i t i v e and s t a t i s t i c a l l y s i g n i f i c a n t r e l a t i o n s h i p between f u l l bole pulp y i e l d and tree s i z e (Figure 39), the slope of the r e l a t i o n s h i p i s small and one can expect l i t t l e r e l a t i v e change i n f u l l bole pulp y i e l d over a f a i r l y wide range of diameter c l a s s e s . This conclusion i s supported by the r e s u l t s presented i n Table 36 which i n d i c a t e that there were not s i g n i f i c a n t d i f f e r e n c e s among diameter classes but that there were s i g n i f i c a n t d i f f e r -ences between trees w i t h i n diameter c l a s s e s . Table 36. Analysis of variance of duplicate determinations of f u l l bole pulp y i e l d for two trees i n each of f i v e diameter classes of 100-year-old lodgepole pine trees. Source df Mean F square Among diam. clas s e s 4 3.73 2.79 Between trees within diam. classes 5 1.34 8.67 Er r o r 10 0.15 To t a l 19 1.22 p.05 s i g n i f i c a n c e l e v e l F s ^ i r ^ 3.33, ^ j S 1 3 5.19 These r e s u l t s generally confirm the r e s u l t s reported by Chidester et a l . , (1939) which indicated that the y i e l d of jack pine pulp varied l i t t l e when considered on a weight b a s i s . Chidester et a l . , (1939) did report, however, that when considered on a volume b a s i s , because of the higher bulk density of slower growing trees, k r a f t pulp y i e l d s were Figure 39. THE RELATIONSHIP B E T W E E N T O T A L U N S C R E E N E D PULP YIELD A N D TREE SIZE. 97 appreciably higher for the smaller sizes than for the larger sizes. Although tree size or growth rate did appear to affect the stength proper-ties of jack pine pulp (Chidester e_t al_. , 1939) , its influence was con-founded by variation resulting from position within the tree and no clear-cut relationships were apparent. Pulp yield would be expected to decrease with tree age because of increased percent juvenile wood (Keays and Hatton, 1971a). There is no obvious reason why pulp yield should decrease with decreasing dbh at the same tree age, unless juvenile wood increases on a weight basis. Variation within trees A considerable amount of research has been devoted to studying within-tree variations in the physical and chemical properties of bole wood. These properties are of particular interest when considering the potential utilization of the non-merchantable top. Based upon a very comprehensive review of these properties, Keays (1971b) presented a general comparison of the physical and chemical properties of the non-merchantable top and merchantable bole for coniferous species (see Table 37). 98 Table 37. General d i f f e r e n c e s i n the wood and pulp c h a r a c t e r i s t i c s of non-merchantable tops compared with merchantable boles for coniferous s p e c i e s . 1 C h a r a c t e r i s t i c Non-merch. top r e l a t i v e to merch. bole Known general wood or f i b r e c h a r a c t e r i s t i c s : S p e c i f i c g r a v i t y Lower Fib r e length Shorter C e l l w all thickness Thinner Wind throw Higher Percentage knots Higher Percentage r e a c t i o n wood Higher Percentage l i g n i n Higher Percentage a l p h a - c e l l u l o s e Lower Pulp c h a r a c t e r i s t i c s ( a n t i c i p a t e d q u a l i t y based on known wood and f i b r e c h a r a c t e r i s t i c s ) : Weight y i e l d Lower Tear f a c t o r Lower Burst f a c t o r Higher T e n s i l e strength Higher Beating time Faster Percent knotter and screen r e j e c t s Higher 1 Source: Keays (1971b) Study of the within-tree v a r i a t i o n s i n pulp y i e l d and q u a l i t y was l i m i t e d to unreplicated tests of pulp from eight d i f f e r e n t l o c a t i o n s w i t h i n one sample tree as out l i n e d on page 77 of t h i s t h e s i s . Table 38 and Figures 40 to 44 i l l u s t r a t e both the v a r i a t i o n w i t h i n the tree and the d i f f e r e n c e s between the merchantable bole and the non-merchantable top. 99 Figure 40. UNSCREENED YIELD (AT 20 PERMANGANATE NUMBER! FOR KRAFT PULP FROM SEVERAL LOCATIONS WITHIN A 100-YEAR-OLD LODGEPOLE PINE TREE. A - Rot torn Quarter B ~ S«cond Quortar C - Third Qvarhrr D - Top Quart** Non - M«r<h. Top 100 Figure 41. BURST FACTOR {AT 300 ml CSF) FOR UNBLEACHED KRAFT PULP FROM SEVERAL LOCATIONS WITHIN A 100-YEAR-OLD LODGEPOLE PINE TREE. A - Bottom Quor te r 8 - Second Quarter C - Third Quarter 0 - Top Quarter N o n " Merch. Top Figure 42. TEAR FACTOR (AT 300 ml CSF) FOR UNBLEACHED KRAFT PULP FROM SEVERAL LOCATIONS WITHIN A 100-YEAR-OLD LODGEPOLE PINE TREE. A - Bottom Quarter B - Second Quart< C - Third Quar te r D - Top Quarter N o n *~March Top Figure 43. BREAKING LENGTH (AT 300 ml CSF) FOR UNBLEACHED KRAFT PULP FROM SEVERAL LOCATIONS WITHIN A 100-YEAR-OLD LODGEPOLE PINE TREE. Figure 44. BULK FACTOR (AT 300m! CSF) FOR UNBLEACHED KRAFT PULP FROM SEVERAL LOCATIONS WITHIN A 1O0-YEAR-OLD LODGEPOLE PINE TREE. S 13000 A - Bottom Quar t** B ~ Second Quarter C - Third Quorter D - Top Quarter A - Bottom Quorter B ~ Second Quarter C - Third Quarter 0 ~ Top Quartet l l N o n - Merch . Top i A B C D , , A B C D | i , i , 1 Merch. Bole N o n - M e r c h . Top 101 Table 38. A comparison of the yield and quality of unbleached kraft pulps from the merchantable bole, non-merchantable top, and f u l l bole of a 100-year-old lodgepole pine tree. Component Adjusted Pulp strength at 300 ml CSF  unscreened yield1 Burst Breaking Tear Bulk (%) factor length (m) factor (cc/gm) Non-merchantable top 46.01 103.0 13,751 98 1.36 Merchantable bole 47.20 104.2 13,473 112 1.41 Full bole 47.00 104.0 13,524 110 1.40 Prorated averages of observations at four different locations in the non-merchantable top and four locations in the merchantable bole. With the exception of burst factor, the results presented in Table 38 were as expected. As can be seen from the results in Table 38, the yield and quality of pulp from the f u l l bole differ l i t t l e from those of the merchantable bole. The large decline in pulp yield observed in the third section of the merchantable bole (Figure 40) was unexpected and no reason for this decline was apparent. A similar result was obtained for a check cook of chips from the same location of the same tree. This result demonstrates the need for careful sample selection when making comparisons of variation both within and between trees. Pulp yields from the non-merchantable top and the merchantable bole were slightly higher than the yields (44.43 and 45.77 percent, respectively) from these components of lodgepole pine grown in Alberta as reported by Szabo and Keays (1973). However, this result was not unexpected because Szabo and Keays (1973) did observe small yield variations from location to location. i 102 Conclusions Some of the methods used in this thesis to determine pulp quality are novel and as such warrant further discussion. The decision to limit the quality tests to pulp from one or two PFI mill runs instead of establishing conventional beater curves was made in order to minimize the amount of sample preparation, cooking time, cooking materials, and paper testing required. For example, one-fifth to two-fifths as much paper testing was required using the PFI mill method as would have been required to establish conventional five-point beater curves for each component of each tree. In addition, the amount of pulp required by the PFI mill method was considerably less than would be required using conventional methods thus simplifying sample collection and facilitating the use of bombs for cooking. This in turn greatly increases the flexibility of the experimental design possible for a given cook. Subsequent tests (Keays, 1972) of the two methods, based on ten replications each using Douglas f i r pulp, indica-ted that the confidence intervals for burst factor, tear factor, breaking length and bulk tests at 300 CSF were 1.02, 3.58, 143.39, and 0.007, respectively for the PFI mill method as opposed to 1.55, 4.52, 337.02, and 0.012, respectively for the conventional five-point beater curve. These results suggest that a higher degree of precision can be obtained by the modified PFI beater procedure, involving 1 or 2 points than can be obtained from a f u l l beater curve (5 points) using a Valley beater. 103 The results of this study indicate that the amount of fibre contained in those components presently considered non-utiliz-able is substantial (Figure 38). Of those presently unutilized com-ponents, the non-merchantable top is of immediate interest because: a. The harvesting and hauling of this component can be easily accommodated by present (full tree) logging practices, and b. This component can be processed (debarked and chipped) by conventional mill equipment. In mature, fully-stocked lodgepole pine stands the number of trees in the 4- to 8-inch diameter classes is substantial (Table 32) and as is demonstrated by the results presented in Figure 38 the percentage of the f u l l bole contained in the non-merchantable top within this diameter range varies from 100.0 to 5.5 percent. It is apparent therefore, that the yield of wood fibre per acre could be greatly increased by harvest-ing to a 1.0-inch top diameter. As can be seen from the results presented for the fu l l bole in Table 38, pulp from this component in combination with pulp from the merchantable bole is of essentially the same quality as pulp from the merchantable bole exclusively. These results suggest therefore that immediate consideration should be given to the utilization of the non-merchantable top in combination with the merchantable bole (i.e., the fu l l bole) because commercial u t i l -ization of this wood resource is dependent upon the economics of harvesting and processing and not pulp yield and quality. The results in Table 35 and Figure 39 which show a positive significant relationship between f u l l bole pulp yield and tree size (which in this study can be equated with growth rate because the trees 104 are essentially even-aged) are of particular interest when considering the feasibility of high-yield silvicultural practices. If forest fertilization, which usually results in a reduction in wood specific gravity and pulp strength characteristics (Sastry et a l . , 1972), were to be practiced on trees genetically selected to give an increase in specific gravity, the results obtained suggest that a substantial increase in yield of fibre per acre per year might be achieved with no change in pulp yield or quality. The branch component was the smallest quantitatively but the most variable of the components examined in this study (Table 31). The results presented in Tables 33 and 34 indicate that the yield and quality of pulp produced from the branch component are substantially less than are those of pulp from the fu l l bole. Commercial ut i l i z a -tion of this component in the foreseeable future is doubtful because of the erratic nature of this component in terms of quantity and because of the inferior yield and quality of branchwood pulp. In addition, i t is doubtful that branches could be successfully debarked and chipped using conventional mill equipment. Small amounts of branchwood pulp might be used in combination with full bole pulp; however, these components would have to be cooked separately using different cooking conditions for each component in order to obtain pulp of equal permanganate numbers (Table 30 and Figure 36). Studies by Keays and Hatton (1971a,b) have suggested that branchwood pulp may be of use as a specialty pulp. 105 As can be seen from the results presented in Table 31 and Figure 38 the root-stump system represents a substantial pro-portion of the total tree (approximately 20 percent). Despite the relatively high yield and quality of pulp obtainable from this com-ponent (Tables 33 and 34), commercial utilization of this component, in the foreseeable future, is doubtful for the following reasons: a. Extraction of the root-stump system would be difficult and time consuming, and, because of the bulky and irreg-ular nature of this component, conventional hauling methods would be unsatisfactory. b. Major problems in removing soil and rock particles from this component (which should be done in the bush prior to hauling in order to minimize transporation costs), and processing (including debarking and chipping) have yet to be overcome. c. In addition, studies of the effect of root extraction on the forest ecosystem with special consideration being devoted to soil erosion, soil water patterns, nutrient cycling, future site productivity, and wildlife have yet to be carried out. In summary therefore, the results presented herein demon-strate that a large proportion of the wood in lodgepole pine trees and stands is not presently utilized. With the exception of branches, these unutilized components can be converted into kraft pulp of a relatively high yield and quality. Immediate consideration should be devoted to the utilization of the non-merchantable top. U t i l i z -ation of the root-stump and branch components is doubtful in the near 106 future. Although further study is warranted, i t appears that cultural practices may be carried out to increase the yield of wood fibre per acre without reducing the yield and quality of pulp. In anticipation of complete-tree utilization in the future, further study should be devoted to determine the effects of this practice on the future productivity of the total forest ecosystem. 107 CHAPTER VI COMPLETE-TREE UTILIZATION OF HIGH STAND DENSITY 100-YEAR-OLD LODGEPOLE PINE Introduction Lodgepole pine often regenerates over-abundantly following w i l d f i r e . Stand d e n s i t i e s of as high as 500,000 stems per acre have been reported i n young stands by Smithers (1959) and even as many as 100,000 l i v i n g stems per acre have been reported i n a 70-year-old stand by Mason (1915). Under these conditions competition and stagnation are severe. Horton (1956) reported a dominant height of only 4 feet i n one 50-year-ol d , dense lodgepole pine stand. Indeed, i t i s u n l i k e l y that stands containing more than 2,000 stems per acre at 90 years of age w i l l ever y i e l d a reasonable merchantable volume (Smithers, 1961). Remedial measures, such as thinning or weeding, may be s u c c e s s f u l l y undertaken i n young stands to overcome stagnation and to increase the y i e l d of usable m a t e r i a l . In older stands such measures hold l i t t l e promise and often the f o r e s t manager i s faced with the expensive p r o p o s i t i o n of c l e a r i n g o f f the e x i s t i n g stand (with l i t t l e or no f i n a n c i a l return) and then p l a n t i n g i f he wants to b r i n g the area back into immediate production. I f the materials removed i n the c l e a r i n g operation could be economically converted i n t o pulp, i t would be p o s s i b l e to defray, at l e a s t i n part, some of the c l e a r i n g and r e f o r e s t a t i o n costs. The purpose of t h i s study i s to describe the y i e l d and q u a l i t y of pulp which could be produced from an overly dense, highly suppressed stand of 100-year-old lodgepole pine t r e e s . 108 Methods and Materials Data c o l l e c t i o n Ten sample trees from Stand 3 (described previously i n Chapter 2) were used to study the pulp y i e l d and q u a l i t y of highly suppressed 100-year-old lodgepole pine trees. No attempt was made to randomize the s e l e c t i o n of the sample trees and only trees apparently free from disease (e.g., dwarf mistletoe and a t r o p e l l i s canker)*were chosen to cover a range of diameter c l a s s e s . The trees were cut at a 1.0-foot stump, and weighed with branches and f o l i a g e i n t a c t ( t o t a l above-ground fresh-weight) and with the branches and f o l i a g e removed ( t o t a l stem fresh-weight). A 1.0-inch thick d i s c , f or moisture content determinations (as previously ou t l i n e d i n Chapter 2) was removed at stump height and at 6.0-foot i n t e r v a l s above stump height. Each tree was divided into three components: root plus stump, bole, and branches plus needles. The branches and needles were drie d (under s i m i l a r conditions to those described previously i n Chapter 2), and because none of the branches were greater than or equal to 1.0-inch i n diameter they were discarded. The oven-dry weights of the bole and root-stump components were determined as previously described i n Chapter 2. The f u l l bole and the root-stump components of each of the ten trees were then transported to the Western Forest Products Laboratory. Subsequently, one of the sample trees was rejected because i t s root-stump component was misplaced i n t r a n s i t . Table 39 presents some of the c h a r a c t e r i s t i c s of the nine suppressed trees used for the pulp study. The e f f e c t s of a t r o p e l l i s canker and dwarf mistletoe on pulp y i e l d and q u a l i t y have been reported by Hunt and Keays (1970), and Hunt (1971), r e s p e c t i v e l y . 109 Table 39. Characteristics of the nine suppressed, 100-year-old lodgepole pine trees used in this study. Tree characteristic Mean Standard Min. Max. dev. value value Diameter (in.) 2.0 0.9 1.2 4.5 Height (ft.) 22.0 5.5 13.2 33.7 Crown length (ft.) 12.5 4.8 8.0 23.7 Crown width (ft.) 3.2 1.0 1.5 4.0 Oven-dry stem weight (lb.) 20.3 16.3 2.1 55.5 Oven-dry root-stump weight (lb.) 8.1 6.3 0.6 22.0 Upon arrival at the laboratory the wood material was stored at a temperature below freezing. Each component was barked by hand. The barked bole sections were chipped in a CAE experimental chipper (courtesy of the British Columbia Institute of Technology) and the barked root-stump components were slabbed on a band saw at a 40°-angle to their grain to give discs 1/2- to 5/8-inch thick, which were reduced to chips by hand. The chips were screened on a Williams chip classifier with 11/8-, 7/8-, 5/8-, and 3/8-inch openings, and a l l oversized chips (retained on the 11/8-inch screen) and fines (chips which passed through the 3/8-inch screen) were discarded. The accept chips were recombined, air dried to a moisture content of about 8 percent, thoroughly mixed, bagged, and stored at 34°F. Preparation and testing of paper from the bole component A random sample of 500 grams of air-dry bole chips was taken from each tree. The chip samples were thoroughly blended together in a laboratory chip mixer (Hatton and Keays, 1970) for 15 minutes. Using the method outlined by Tappi Standard T18 os-53 (method 2), the specific gravity of the chip mixture was determined to be 0.40. Prior to cooking 5 chip 110 moisture determinations were made. The chips were pulped by the k r a f t process using the pulping conditions presented i n Table 40. Table 40. Kraft cooking conditions f o r the bole chip mixture of nine suppressed, 100-year-old lodgepole pine trees. S u l f i d i t y (%) 25.0 E f f e c t i v e a l k a l i (%) 17.0 Time to max. temp. (min.) 135.0 Time at max. temp. (min.) 80.0 Max. temp. (°C) 170.0 Liquor-to-wood r a t i o 4:1 The preceding pulping conditions (Table 40) were selected to obtain an unbleached pulp permanganate number of close to 20. The laboratory pulping equipment used i n t h i s study was a s t a i n l e s s s t e e l Weverk research digester with a 1.0-cubic foot capacity. Upon completion of the cooking the pulp was thoroughly washed, centrifuged for 5 minutes, and mixed. Five separate oven-dry a l i q u o t s of pulp were used f o r y i e l d determinations p r i o r to screening. The amount of r e j e c t s was determined by weighing the oven-dry material retained on a 10-cut p l a t e of a Va l l e y laboratory f l a t screen. Unbleached permanganate and Kappa numbers were determined on the screened pulp following Tappi Standards T214 ts-50 and T236/m-60, r e s p e c t i v e l y . Three determinations of each were made on the pulp. One beater run was done for the pulp using a Va l l e y laboratory beater f o l l o w i n g the method outlined by Tappi Standard T200 ts-66. Pulp freeness was tested (Tappi Standard T227 Os-58) and pulp samples f o r handsheets were c o l l e c t e d at time i n t e r v a l s of 0,5,25,50,60 and 72 minutes. Six handsheets f or each of the previously mentioned beating times were prepared according to Tappi Standard T205 m-58. The f i v e best hand-I l l sheets for each beating time were tested for bursting and tearing strengths and breaking length following Tappi Standard T220 m-60. Preparation and testing of paper from the root-stump component component of the trees was small i t was impossible to follow the preceding procedures used for the bole component and consequently the following procedures were used. A random sample of 50 air-dry grams of root-stump chips was taken for each tree. The chip samples were thoroughly blended together in a laboratory mixer for 15 minutes. Duplicate moisture content determinations of the 60 air-dry grams of randomly selected chips from the mixture were obtained. Four, 60-gram (a.d.) chip samples, obtained at random from the mixture, were sealed in stainless steel bombs (each of 735-ml capacity) and placed in the research digester. In order to obtain an unbleached pulp permanganate number of close to 20, the root-stump chips were pulped by the Kraft process using the pulping conditions presented in Table 41. Because the amount of chips available from the root-stump Table 41. Kraft cooking conditions for the root-stump chip mixture of nine suppressed, 100-year-old lodgepole pine trees. Sulfidity Effective alkali Time to max. temp. Time at max. temp. Max. temp. Liquor-to-wood ratio (%) (%) 25.4 14.8 135 (min.) (min.) (°C) 75 170 4.5:1 The cooked chips from each bomb were disintegrated for 5 minutes and the resulting pulp was thoroughly washed. Two of the four samples, to 112 be used for screenings, permanganate number, and y i e l d determinations, were oven-dried to a constant dry-weight for a minimum of 16 hours at 105°C. The remaining two samples, to be used f o r PFI m i l l and strength determinations, were a i r - d r i e d f o r a s i m i l a r time period. The pulp from each sample was then r e d i s i n t e g r a t e d f o r 5 minutes and the amount of r e j e c t s was determined by weighing the oven-dry material retained on a 10-cut plate of a V a l l e y laboratory f l a t screen. The screened pulp from each bomb was centrifuged for f i v e minutes, thoroughly mixed, and conditioned to a uniform moisture content of about 8 percent. Unbleached permanganate numbers were determined on the screened pulp used for the y i e l d determinations following Tappi Standard T214 ts-50 (using 40 ml of 0.1 N KMn0i») . The screened pulp from the two samples to be used f o r strength determinations were thoroughly mixed. This mixture was divided into two equal aliquots from each of which a sample of 24.0 grams (o.d.) was withdrawn. Each sample was d i s i n t e g r a t e d at a consistency of approximately 1.2 percent u n t i l free from f i b r e bundles. Each sample was then adjusted to a consistency of 10 percent and placed i n the PFI m i l l . A pulp freeness of 306 CSF was obtained at 22,000 beating revolutions of the PFI m i l l . For the reasons stated previously, t h i s was considered to be s u f f i c i e n t l y c l o s e to the target freeness of 300 CSF and consequently the second PFI m i l l run was not c a r r i e d out. Pulp freeness was tested following Tappi Standard T227 os-58 and s i x handsheets were prepared according to Tappi Standard T205 m-58. The f i v e best handsheets were tested for breaking length, and burst and t e a r i n g strength following Tappi Standard T220 m-60. 113 Analysis Because, for the reasons outlined in Chapter 5, i t is desirable to compare pulp yields at a uniform permanganate number, an adjusted pulp yield at permanganate number 20 was calculated using the following formula: obs obs where: Y20 = yield at permanganate number 20 Y , = observed yield obs J K , = observed permanganate number obs & B = 0.38 for bole pulp = 0.36 for root-stump pulp. Results and Discussion In the following section comparisons are made between the yield and quality of pulp produced from the bole and root-stump components of the suppressed trees, and the average yield and quality of pulp produced from more normally grown lodgepole pine trees observed in Chapter 5. For the bole components a further comparison is made with the results reported for lodgepole pine by Hatton and Keays (1970) which were obtained under similar laboratory conditions (i.e., the same equipment, test procedures and personnel), from merchantable trees of the same age as those used in the present study. Table 42 presents unadjusted and adjusted (to permanganate number 20) unscreened pulp yield data obtained for the suppressed trees used in this study in comparison with similar data obtained for normally grown lodgepole pine and for the reference standard. 114 Table 42. A comparison between unscreened pulp y i e l d data of suppressed pine trees, more normally grown pine trees and the reference standard. Unadjusted Permanganate Adjusted Kappa Screenings y i e l d (%) number y i e l d (%) no. (%) Bole: Supp.pine: mean 44.17 17.9 44.96 25.9 0.60 range 43.88-44.36 44.67-45.15 Normal range 45.54-48.86 18.8-23.3 45.33-48.78 0.11-2.20 pine: 1* mean 47.11 20.3 46.99 0.84 Ref.stand: mean 43.70 1 17.9 1 44.8 2 0.40 3 range 43.65-43.761 17.85-17.90 Root-stump: Supp.pine: mean 44.86 22.0 44.14 0.63 range 44.40-45.31 21.4-22.6 43.90-44.37 0.20-1.06 Normal pine: 1* mean 46.00 19.8 46.06 0.64 range 41.26-47.46 18.6-21.1 41.76-47.50 0.16-1.78 1 Ref. stand, from Table I I I , Cook 75, Hatton and Keays (1970). 2 Ref. stand, from Table VI, Hatton and Keays (1970). 3 Ref. stand, from Cook 75, unpublished data (Hatton, 1972). "* Table 33, Chapter 5 of t h i s t h e s i s . Table 43 presents the beating times,burst f a c t o r s , tear f a c t o r s , and breaking lengths at 500 and 300 ml CSF of the pulp handsheets prepared from the suppressed trees i n comparison with data obtained f o r more normally grown pine and f o r the reference standard. 115 Table 43. A comparison between the mean unbleached pulp quality of the suppressed pine trees, more normally grown pine trees and the reference standard. 500 ml CSF 300 ml CSF Burst Tear Break, factor factor length (m) Burst factor Tear factor Break, length (m) Bole: Supp.pine 94 Normal pine1 — Ref. Stand.2 100±3 Root-stump: Supp.pine Normal pine1  103 114 ±9 12,750 13,800±400 107 97 106±1 72.4 74.6 93 114 105±4 131.0 116.4 14,150 12,598 14,400±350 9,761 9,887 1 Table 34, Chapter 5 of this thesis. 2 Keays (1972). As demonstrated by the results presented in Table 42, the un-screened yield of pulp produced from the suppressed trees is essentially the same as that obtained for more normally grown lodgepole pine. Similarly, no substantial difference was noted in the percent screenings. The results also indicate that the quality of unbleached pulp produced from the sup-pressed trees is comparable to that produced from normally grown trees (Table 43). For the full bole component the suppressed pine tended to develop their strength more rapidly. Conclusions No data are presently available for the acreage of high density lodgepole pine stands similar to the one examined in this study. However, as is demonstrated by the results presented in Table 32, even more normally 116 grown stands of 100-year-old lodgepole pine may contain a large number of small trees s i m i l a r to those examined i n t h i s study. The y i e l d and q u a l i t y of pulp produced from these suppressed trees was e s s e n t i a l l y the same as that produced from more normally grown trees (Tables 42 and 43). These r e s u l t s suggest therefore that commercial u t i l i z a t i o n of t h i s wood resource i s dependent upon the economics of harvesting and processing and not pulp y i e l d and q u a l i t y . The r e s u l t s presented herein suggest that the quantity of f i b r e (Table 7), and q u a l i t y and y i e l d of pulp from these overly dense stands may be s u f f i c i e n t to defray the costs of returning such areas to f u l l production. Although u t i l i z a t i o n of t h i s resource i n the foreseeable future i s doubtful, some p o t e n t i a l advantages i n i t s u t i l i z a t i o n include: a. The small s i z e of t h i s material enhances the prospects of mechanized hauling and handling ( i . e . , bales) and harvesting ( i . e . , a ' f i b r e combine 1 such as that reported on i n the B r i t i s h Columbia Lumberman (1971)). However, because the long bast f i b r e s found i n the bark of deciduous species (Crossley, 1938) are absent i n the bark of coniferous species, bark removal p r i o r to cooking may complicate the development of a ' f i b r e combine' s u i t a b l e for use with coniferous species. Research on improved methods f o r separating bark from chips i s proceeding. b. U t i l i z a t i o n of dense stands could return land to a more pro-ductive state and t h i s aspect might f a c i l i t a t e negotiations f o r reduced stumpage charges or g r a n t s - i n - a i d of improved management. c. Present boundaries between commercial and non-commercial forests might be revised s u b s t a n t i a l l y , because s i t e s presently consid-ered incapable of growing trees to merchantable s i z e s might become operable. 117 CHAPTER VII SUMMARY AND SUGGESTIONS FOR FURTHER RESEARCH The objective of this thesis is to present the results of studies of dry-matter production, tree growth, and complete-tree utilization of 100-year-old lodgepole pine based on data obtained from the intensive examination of some forest-grown trees from southwestern Alberta. The results are used to estimate and compare the total standing crop of 100-year-old lodgepole stands grown over a range of site and stand density conditions throughout Alberta. In addition, comparisons are made between the above-ground dry-matter production of young lodgepole pine and Populus stands in Alberta. Chapter 2 of this thesis presents allometric relationships for the estimation of component dry-weights for 100-year-old lodgepole pine trees grown in average and high density stands. Errors resulting from the use of logarithmic equations are examined and factors to correct these errors are presented. With the exception of the crown components, these errors are negligible. First approximations of the total standing crop of 100-year-old lodgepole pine stands, by components, were estimated for a range of site and stand density conditions in Alberta. The combined variable of stand basal area times mean stand height appears to provide reliable estimates of total standing crop and component biomass. Component biomass was found to be inversely related to number of stems on a given site. The relationships and approximations presented in Chapter 2 lack generality because of the limited age, site and stand conditions examined. This limitation can be rectified in the future by a more extensive exam-118 ination of these various conditions. Chapter 3 of this thesis presents a comparison of the above-ground organic matter production of lodgepole pine and Populus stands of similar ages grown on similar sites. For the sites examined the above-ground standing crop of the pine stands were substantially greater than the above-ground standing crop of the Populus stands. Further comparisons of organic matter production between species over a range of age, site and stand density conditions would be useful in determining the maximum productivity and the suitability of a given species on a given site. The relationships between several crown parameters and radial, cross-sectional area and section volume growth of 100-year-old lodgepole pine trees were examined in Chapter 4. Although tree volume growth was closely related to the dry weight of needles of the tree, no relationship was found between the efficiency of production (volume growth per unit foliage) and tree size as had previously been found for some species. Thinning did not appear to affect the pattern of growth within trees. Although some relationships were established between the amount of growth at a given point within a tree and the amount of foliage at or above that point, these relationships were not as strong as had been observed previously for other species, probably because of the advanced age of the sample trees. Further study of the patterns of growth within and between trees should be devoted to younger, more vigourously growing trees with consideration being given to data analysis prior to sample collection. Chapters 5 and 6 of the thesis are devoted to the complete-tree utilization of 100-year-old lodgepole pine trees grown in average and high density stands. Relationships were developed to establish the oven-dry, bark-free quantity of the various components expressed as a percentage of 119 the oven-dry, bark-free f u l l bole (1.0-inch top). Because only ten trees were used i n t h i s study, further development of these r e l a t i o n s h i p s i s warranted. K r a f t pulp y i e l d and q u a l i t y data were obtained for each com-ponent and r e l a t e d to tree s i z e . The r e s u l t s indicated that f a s t e r tree growth r e s u l t s i n a s l i g h t l y higher y i e l d of pulp from the f u l l bole with no apparent a f f e c t on pulp q u a l i t y . Tree s i z e and growth rate were not s i g n i -f i c a n t l y r e l a t e d to the y i e l d and q u a l i t y of pulp from the other components. Because the quantity of wood f i b r e contained i n the non-merchantable top (4.0-inch to 1.0-inch top) of trees 4.0 to 8.0 inches i n diameter, which may make up a s u b s t a n t i a l proportion of mature lodgepole pine stands, i s l a r g e , and because the y i e l d and q u a l i t y of pulp from t h i s component i s high, immed-i a t e consideration should be given to the u t i l i z a t i o n of t h i s component. Although the y i e l d and q u a l i t y of pulp from the root-stump system are r e l a t -i v e l y high, u t i l i z a t i o n of t h i s component i n the near future i s doubtful because of the t e c h n i c a l problems associated with e x t r a c t i n g , cleaning, transporting, and processing t h i s component. The r e l a t i v e l y low y i e l d and q u a l i t y of pulp from branchwood coupled with the problems of processing t h i s component suggest that imminent u t i l i z a t i o n of lodgepole pine branches i s doubtful. The y i e l d and q u a l i t y of pulp from highly suppressed 100-year-old pine trees were e s s e n t i a l l y equal to the y i e l d and q u a l i t y of pulp from mature lodgepole pine. The r e s u l t s presented h e r e i n i n no way attempt to examine the e f f e c t s of complete-tree u t i l i z a t i o n on the f o r e s t ecosystem. Future study of n u t r i t i o n a l make-up of the various components of trees growing under various f o r e s t conditions, s i m i l a r to that reported by Boyle and Ek (1972), i s necessary to e s t a b l i s h the degree to which complete-tree u t i l i z a t i o n taxes the forest s i t e . In a d d i t i o n , study of the e f f e c t s of complete-tree u t i l i z a t i o n on s o i l erosion, s o i l moisture patterns and w i l d l i f e management 120 are warranted. 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Challenge of complete tree u t i l i z a t i o n . Forest Prod. J . 18 (4): 83-86. 131 , and J. P. Kramer. 1952. The effect of pruning on the height and diameter growth of loblolly pine. J. Forest. 5_0 (6):474-479. , L. Strand, and R. A. Altenberger. 1964. Preliminary fresh and dry weight tables for seven tree species in Maine. Maine Agr. Exp. Sta., Tech. Bull. 12. 76 pp. , and A. J. Chase. 1965. Fiber weight and pulping characteris-tics of the logging residue of seven tree species in Maine. Maine Agr. Exp. Sta., Tech. Bull. 17. 44 pp. , P. N. Carpenter, and R. A. Altenberger. 1965a. Preliminary tables of some chemical elements in seven tree species in Maine. Maine Agr. Exp. Sta., Tech. Bull. 20. 88 pp. , L. Hoar, and M. Ashley. 1965b. Weight of wood substance for components of seven tree species. Tappi 48_ (8) : 466-469.. , and V. P. Guinn. 1966. Chemical elements in complete mature trees of seven species in Maine. Tappi 49_ (5): 190-197. , and P. M. Carpenter. 1967. Weight, nutrient elements and productivity studies of seedlings and saplings of eight tree species in natural ecosystems. Univ. Maine, Maine Agr. Exp. Sta., Tech. Bull. 28. 39 pp. Zar, J. H. 1968. Calculation and miscalculation of the allometric equation as a model in biological data. Bio. Sci. IB (12) : 1118-1120. 132 APPENDIX I. Lesser vegetation present i n Stands 1 and 2. Aster conspicuus L i n d l . Cornus canadensis L. Cornus s t o l o n i f e r a Michx. Elymus innovatus Beal. Epilobium angustifolium L. Equisetum hyemale L. Galium labradoricum Wieg. Lathyrus ochroleuvus Hook. Linnaea b o r e a l i s L. var. americana (Forbes) Rehd. Lonicera d i o i c a L. var. glaucenscens (Rydb.) Butters Petasites palmatus (Ait.) A. Gray Populus tremuloides Michx. Pyrola a s a r i f o l i a Michx. Rosa a c i c u l a r i s L i n d l . S m i l i c i n a s t e l l a t a (L.) Desf. Streptopus amplexifolius (L.) DC. Thalictrum occidentale A. Gray Viburnum trilobum Marsh. V i o l a rugulosa Greene Showy aster Bunchberry Red o s i e r dogwood Hairy wild rye Fireweed Scouring rush Bedstraw Pea vine Twin-flower Twining honeysuckle Palmate-leaved c o l t s f o o t Trembling aspen Common pink wintergreen P r i c k l y rose Star-flowered Solomon's-Seal Twisted-stalk Western meadowrue High-bush cranberry Western Canada v i o l e t 133 Appendix 1-1. DRY TOTAL TREE WEK5HT-DBH ALLOMETRY Appendix H . - 2 . 0 R Y ABOVE-GROUND WE1GHT-DBH ALLOMETRY OF MATURE WHITE SPRUCE. OF MATURE WHITE SPRUCE. O«NI«,.I M H I * . I Appendix B - 3 . DRY STEM WEIGHT-BASAL AREA RELATIONSHIP OF MATURE WHITE SPRUCE. Appendix 1 - 4 . DRY MERCHANTABLE STEM WEIGHT-BASAL AREA RELATIONSHIP OF MATURE WHITE SPRUCE. •* * A - i t * IX M • •» .4 .« Jk 10 I J u I M e r a l ATM [«lt] TrM total A/*o(iQ.ft.) 134 A p p e n d i x B-5. D R V N E E D I E W E I G H T - D B H A L L O M E T R Y OF M A T U R E W H I T E S P R U C E . A p p e n d i x 1 - 6 . D R Y B R A N C H W E I G H T - D B H A L L O M E T R Y OF M A T U R E W H I T E S P R U C E . / I / 1 IW*2.030 Log^O- 0.163 t "0.9 V.* i n - IB M.2 lb, [24 8%] • / • / / . ' •or: 5 n • 18 '» lb. [4141b] 097 \ 08H (MV) A p p e n d i x J T - 7 . D R Y R O O T - S T U M P W E I G H T - B A S A L A R E A R E L A T I O N S H I P O F M A T U R E W H I T E S P R U C E . I i £ to M + SW- 215.78 8A-I2.50 r* • 0.988 n • 18 S..,' 11.7 lb. [14.9%] TtM towl AfM(tq.fl.) 135 APPENDIX III. Radial growth and foliage weight distribution diagrams of twenty, 100-year-old lodgepole pine trees. i Legend: 5-yr. radial growth (mm) <J > Foi. wt. (lb.) 2 f t . above «j > Cumulative f o i . wt. (lb.) above < J - b, 136 TREE I TREE 2 abbot- 6.6 !•. Wight • 63.3 ll. . JO-S' growth (mm) - 30 Dry woighl of (Oliogo Hb) 5-yoor rodiol growth Lmn) 3 10 15 Dry woigta of loliogo (IbJ dbhob • 6.3 i«. Wight • 34.B fl T R E E 3 dbhab • 3.3 in. •night • 34.3 ft. K> 3 3-jroor radial growth \mm) 3 K) Dry woighl of •ollogo lib} 3-yoor radial growth 1»n) Ory woight of loliogo (IbJ T R E E 4 «bhob- 3.] i„. hoight • 43.3 (i. 137 TREE 5 4bkob • K.Sr>. hoigM • 67.T f«. 5 it) S 20 25 30 35 40 Dry woight of fotiog* fIbJ TREE'6 tfbhob • 7.0 «. bolglit • oJ.« ft. E h 10 J 3-yoor rao*iol growth (mm) 5 to 13 Dry w«ig>t of foliogo (IW T R E E 7 ObHob • 7.0 in. Wight - 34.0 h. J-yOOr rotf%ol Dry w«.gM of foliogo (IbJ T R E E 8 3.8 io. 39.3 ft. 3-yoor rodiol ffrowtn (mm) Dry woighl of foliogo ClbJ 138 JO 40 2 X JO-»• 0 70 to JO 10 to 30 3-yoor radio) Dry Might of foliogo (IbJ 5-yoor rodiol growth (mm) Dry woight ef foliogo (lb) TREE II TREE 12 dbhob • 7.4 ir bight • tl} Dry woight Of foliogo (lb.) 3- Jo 1 dbhob < W.2 io. hoight ' 7J.J ft. » J 3-yoor rodiol growth (»•) S » U 30 2J Dry woight of foliogo (lb) 139 i » 5 3-ytrar rodiol Srowril (mm) TREE 13 dbhob • 11.* h. MgM • 77.4 ft ~3 » 3 55 5 3 5 3 I J -Dty woight of foliogo Ob) TREE 14 • b o o b • 7.2 ov b i g h t • «0.7 K 10 3 5 K U JO 3- poor todiot Dry wMgb* of foiiogo (lb) f j rowrb(M) TREE 15 TREE 16 dbhob • 7 3 io. hoight • 39.1 l l . n i 3-yoor rodiol Kfmak) J 10 13 Dry woighl of foliogo (lb) HZ w 1 3-yoor rodiol growth (oia) dbhob . 7.J i « . hoight • 57J J l . 3 10 13 Dry woighl of foliogo Ob) 140 •0 50 40 s I »• 30 10-0 70 30-40-2 I * JO-2 10-T R E E 17 T R E E 18 dbhob • S3 r». k.igr» • 61.7 It. .bkob : 4.8 in. bight • 47 2 It. K 5 5-yMr rodiol growth iiorn) o 10 3 10 13 Dry weight of foliog. (lb) » 5 5-ywor rodiol growth (mnu 3 n Dry w.ight of foliog. (lb) TREE 19 dbhob • 8.8 in. hoight • 69.2 ft. 3 10 13 20 13 Dry w.ight of foliogo (lb) TREE 20 dbhob • 4.7 in. hoight - 36.2 ft. 

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