UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

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

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Item Metadata

Download

Media
831-UBC_1973_A1 J63_5.pdf [ 7.5MB ]
Metadata
JSON: 831-1.0101020.json
JSON-LD: 831-1.0101020-ld.json
RDF/XML (Pretty): 831-1.0101020-rdf.xml
RDF/JSON: 831-1.0101020-rdf.json
Turtle: 831-1.0101020-turtle.txt
N-Triples: 831-1.0101020-rdf-ntriples.txt
Original Record: 831-1.0101020-source.json
Full Text
831-1.0101020-fulltext.txt
Citation
831-1.0101020.ris

Full Text

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 o f B r i t i s h Columbia, 1966 M.F., U n i v e r s i t y o f 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 a c c e p t t h i s  t h e s i s as. c o n f o r m i n g t o the r e q u i r e d  THE UNIVERSITY OF BRITISH COLUMBIA A p r i l , 1973  standard  In p r e s e n t i n g an  this  thesis i n partial  advanced degree a t t h e U n i v e r s i t y  the  Library  s h a l l make i t f r e e l y  f u l f i l m e n t of the requirements f o r o f B r i t i s h Columbia, I agree  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 extensive for  copying of t h i s  thesis  s c h o l a r l y p u r p o s e s may b e g r a n t e d by t h e Head o f my D e p a r t m e n t 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 understood that  of t h i s  thesis f o rfinancial  written  permission.  Department o f  April.  gain  Forestry  The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, C a n a d a  Date  that  1973  Columbia  copying or p u b l i c a t i o 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  i ABSTRACT Chairman: Dr. D. D. Munro  Lodgepole pine (Pinus contorta Dougl. var. l a t 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,  c.  and  to compare the above-ground total standing crops of similarly aged lodgepole pine and Populus stands grown on similar s i t e s . 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 i n 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.  ii 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 dryweights 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 s i t e . Fresh- and dry-weight estimates, by component, were obtained from two paired, young lodgepole pine and Populus stands grown i n 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 i n productivity are discussed. Within and between tree variations i n radial, cross-sectional area, and volume growth were examined for twenty, 100-year-old lodgepole pine trees.  Ten trees were sampled from both a thinned and an  unthinned stand. Although volume growth was found to be highly correlated 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 i n the tree to the needle biomass at or above that point i n the tree are presented.  A large amount of  within tree variation i n growth was not accounted for by these relationships.  Thinning did not appear to affect the pattern of growth of the  trees examined. Suggestions for further research on the growth patterns  iii  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 i n 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 f u 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 i n 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 i n some stands, immediate consideration should be given to i t s u t i l i z a t i o n . Although the yield and quality of pulp from the root-stump system was found to be high, utilization of this component i n the near future i s doubtful because of the technical problems associated with extraction, cleaning, transporation, and processing.  Similarly, utilization of  branches i n the near future i s doubtful because of the low yield and quality of pulp from branchwood and because of the processing problems associated with i t s u t i l i z a t i o n .  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 c r i t i c a l 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 s t a f f , 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 u t i l i z a t i o n 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.  vii TABLE OF CONTENTS Page ABSTRACT  i  ACKNOWLEDGEMENTS  v  TABLE OF CONTENTS  vii  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 Analysis Results and Discussion Component dry-weights of 100-year-old lodgepole pine trees  5 11 16 16  Component dry-weights of 100-year-old lodgepole pine stands Conclusions  28 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 CHAPTER IV VARIATIONS IN STEM GROWTH AS RELATED TO SEVERAL CROWN CHARACTERISTICS OF 100YEAR-OLD LODGEPOLE PINE TREES  54  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 The quantity of each component potentially available from the complete-tree utilization of 100-year-old lodgepole pine trees  87  87  The quality of pulp obtained from the completetree utilization of 100-year-old lodgepole pine trees 91  TABLE OF CONTENTS  (Continued) Page  V a r i a t i o n among t r e e s  91  V a r i a t i o n within trees  97 102  Conclusions CHAPTER V I  COMPLETE-TREE UTILIZATION OF HIGH STAND DENSITY 100-YEAR-OLD LODGEPOLE PINE.....  107  Introduction  107  Methods and M a t e r i a l s  108 108  Data c o l l e c t i o n P r e p a r a t i o n and t e s t i n g o f paper  109  from t h e b o l e component P r e p a r a t i o n and t e s t i n g o f paper from the root-stump  component..  I l l  Analysis..  113  R e s u l t s and D i s c u s s i o n  113  Conclusions  115  CHAPTER V I I  SUMMARY AND SUGGESTIONS FOR FURTHER RESEARCH  REFERENCES CITED  117 121  APPENDICES: I  L e s s e r v e g e t a t i o n p r e s e n t i n Stands 1 and 2 132  I I - l Dry t o t a l t r e e weight - DBH a l l o m e t r y o f mature w h i t e s p r u c e II-2 Dry above-ground weight - DBH a l l o m e t r y o f mature w h i t e s p r u c e  133 133  II-3 D r y stem weight - b a s a l a r e a r e l a t i o n s h i p o f mature w h i t e s p r u c e 133 II-4 Dry merchantable stem weight - b a s a l a r e a r e l a t i o n s h i p o f mature w h i t e s p r u c e  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 1 2 3 4 5  6 7  8  Page Characteristics of Stands 1 and 2 i n 1938 before and after thinning...  6  Characteristics of the three stands of 100-year-old lodgepole pine .  6  Statistical characteristics of the independent variables for 100-year-old lodgepole pine trees  16  Statistical characteristics of the dependent variables for 100-year-old lodgepole pine trees  17  Simple correlation coefficients (r) between tree and component dry-weights and several tree characteristics for 100-year-old lodgepole pine trees  18  Dry-weight allometric relationships for 100-yearold lodgepole pine trees  20  Dry weight of tree components and total standing crop for three stands of 100-year-old lodgepole pine  27  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-yearold lodgepole pine stands  35  Characteristics of the four lodgepole pine and Populus stands  41  Statistical characteristics of the independent variables from twenty, young lodgepole pine trees  45  Statistical characteristics of the dry-weight dependent variables from twenty, young lodgepole pine trees  45  12 13 14  xii LIST OF TABLES (Continued) Table 15  16  17  18  19  20  21 22 23  24  Page Simple correlation coefficients (r) between component dry-weights and several tree characteristics of twenty, young lodgepole pine trees  46  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  Statistical characteristics of the fresh-weight dependent variables from twenty, young lodgepole pine trees  49  Simple correlation coefficients (r) between component fresh-weights and several tree characteristics of twenty, young lodgepole pine trees  49  A comparison of measured fresh weights (lb./ac.) of paired lodgepole pine and Populus stands of similar ages on similar sites  52  A comparison of estimated above-ground dry weight (lb./ac.) of living components i n paired lodgepole pine and Populus stands of similar ages on similar sites  51  Proportion of total above-ground dry-weight by components of major species in study areas  54  Some characteristics of the twenty, 100-year-old lodgepole pine sample trees  62  Simple correlation coefficients (r) between several tree characteristics of the twenty, 100-year-old lodgepole pine sample trees  63  Analyses of variance of radial, cross-sectional area, and volume growth measured at 5.positions within the crown and 5 positions within the clearbole of ten, 100-year-old lodgepole pine trees grown in a thinned stand and ten, 100-year-old lodgepole pine trees grown i n an unthinned stand  66  i  xiii LIST OF TABLES (Continued) Table 25  26  27  28 29 30  31  32  33 34 35  36  Page Simple correlation coefficients (r) between 358 measurements of radial, cross-sectional area and section volume growth, and several tree and section characteristics  . 67  Simple correlation coefficients (r) between 245 measurements of r a d i a l , cross-sectional area and section volume growth i n crown-formed wood, and several tree and section characteristics...  69  Comparison of relative values of kraft pulp from conifer tree components expressed i n terms of the bole  75  Characteristics of the selected 100-year-old lodgepole pine trees...  79  Kraft cooking conditions used for the exploratory cooks  81  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  Bark-free, oven-dry weights of the 100-year-old lodgepole pine tree components as a percentage of f u l l boles  87  Average number of stems per acre i n diameter classes less than 9.0 inches dbhob from eighty-eight stands of 100-year-old lodgepole pine  90  Component pulp yield data from kraft pulping for ten, 100-year-old lodgepole pine trees  92  Component pulp quality at 300 ml CSF from kraft pulping for ten, 100-year-old lodgepole pine trees...  93  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  Analysis of variance of duplicate determinations of f u l l bole pulp yield for two trees i n each of five diameter classes of 100-year-old lodgepole pine trees  95  xiv LIST OF TABLES (Continued) Table 37  38  39  40  41  42  43  Page General differences i n the wood and pulp characteristics of non-merchantable tops compared with merchantable boles for coniferous species  98  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  Characteristics of the nine suppressed, 100-yearold lodgepole pine trees used in this pulp study  109  Kraft cooking conditions for the bole chip mixture of nine suppressed, 100-year-old lodgepole pine trees  110  Kraft cooking conditions for the root-stump chip mixture of nine suppressed, 100-year-bld lodgepole pine trees  Ill  A comparison between unscreened pulp yield data of suppressed pine trees, more normally grown pine trees and the reference standard  114  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 1 2 3  4 5  6 7  8 9 10 11  12 13 14  15  Page  The relationship between moisture content and height i n the tree  8  The relationship between specific gravity and height i n the tree  8  The relationship between bias correction factors and standard errors of estimate (a) of logarithmic equations  15  Total tree weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine  21  Total above-ground component weight - D2H allometry of 'average stand density' 100year-old lodgepole pine  21  Stem weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine  21  Merchantable stem weight - D2H relationship of 'average stand density' 100-year-old lodgepole pine  21  Stem bark weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine  22  Needle weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine  22  Branch weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine  22  Stump plus root weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine  22  Root weight - D2H allometry of 'average stand density' 100-year-old lodgepole pine  23  Total tree weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine  24  Total above-ground component weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine  24  Stem weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine  24  xyi LIST OF FIGURES (Continued) Figure 16 17 18 19  20  21  22  23  24  25  26  Page  Needle weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine  24  Branch weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine  25  Stump plus root weight - D2H allometry of 'high stand density' 100-year-old lodgepole pine  25  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  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  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  The relationship between estimated dry aboveground weight per acre and stand basal area times mean stand height for stands of 100-yearold lodgepole pine  31  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  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  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  The relationship between stem dry-weight and tree diameter of young lodgepole pine trees  48  xvii LIST OF FIGURES (Continued) Figure 27  28  29  30  31  32  33  34  35  36  37  Page  The relationship between branch dryweight and tree diameter of young lodgepole pine trees  48  The relationship between needle dryweight and tree diameter of young lodgepole pine trees  48  The relationship between above-ground dryweight and tree diameter of young lodgepole pine trees  48  The relationship between stem freshweight and D2H of young lodgepole pine trees  50  The relationship between crown freshweight and D2H of young lodgepole pine trees  50  The relationship between above-ground fresh-weight and D2H of young lodgepole pine trees  50  The relationship between needle dryweight and tree diameter for the twenty, 100-year-old lodgepole pine trees  64  The relationship between past 5-year volume growth and present needle dry-weight of twenty, 100-year-old lodgepole pine trees  64  The relationship between volume growth efficiency (cu.ft./lb. dry needles) and tree diameter of twenty, 100-year-old lodgepole pine trees  64  The relationship between unbleached pulp permanganate number and effective a l k a l i for three components of 100-year-old lodgepole pine  82  Relationship between unscreened yield and permanganate number for kraft pulp from three components of 100-year-old lodgepole pine trees...  86  xviii  LIST OF FIGURES (Continued) Figure 38  39 40  41  42  43  44  Page  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  The relationship between total unscreened pulp yield and tree size  96  Unscreened yield (at 20 permanganate number) for kraft pulp from several locations within a 100-year-old lodgepole pine tree  99  Burst Factor (at 300 ml CSF) for unbleached kraft pulp from several locations within a 100-year-old lodgepole pine tree  100  Tear factor (at 300 ml CSF) for unbleached kraft pulp from several locations within a 100-year-old lodgepole pine tree  100  Breaking length (at 300 ml CSF) for unbleached kraft pulp from several locations within a 100-year-old lodgepole pine tree  100  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 i n 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 i s 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 ecosystems 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  f e r t i l i z a t i o n and nutrient flow (Ovington and Madgwick, 1959; Orman and W i l l , 1960; W i l l , 1964; Young et a l . , 1964; Rennie, 1955;  2 Bazilevic and Rodin, 1966, Marchenko, 1967; Duvigneaud and DenaeyerDe 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, i s the primary source of organic matter and potential energy upon which a l l l i f e (with the exception of some bacteria) i s 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 i s 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 i n 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 i n 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 i n 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) i s the best documented, multi-disciplinary ecosystem study available to date. The objective of this thesis i s 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. l a 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 f i r s t 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 mariana (L.)  Canada biomass r e s u l t s have been r e p o r t e d f o r P i c e a  ( M i l l . ) BSP (Weetinan and H a r l a n d , 1964), A b i e s balsamea  Mill.  ( B a s k e r v i l l e , 1965a,b, 1966; Honer, 1971), P i n u s c o n t o r t a  D o u g l . v a r . l a t i f o l i a Engelm. ( K i i l , 1967,  1965; Muraro, 1966; J o h n s t o n e ,  1970a,b, 1971), P i n u s r e s i n o s a A i t . ( S t i e l l ,  1966),  Populus  t r e m u l o i d e s Michx. ( B e l l a and J a r v i s , 1967; B e l l a , 1968, 1970; P e t e r s o n e t a l . , 1970; P o l l a r d , 1972),  P i c e a g l a u c a (Moench) Voss  (Keays and H a t t o n , 1971b), P i n u s b a n k s i a n a Lamb. ( H e g y i , 1972), and Tsuga h e t e r o p h y l l a (Raf.) S a r g . , Pseudotsuga  menziesii  (Mirb.)  F r a n c o , and T h u j a p l i c a t a Donn (Osborn, 1968; Kurucz, 1969; P a i l l e , 1970;  Keays and H a t t o n , 1971a; Smith, 1971; McGreavy, The purpose  1972).  o f t h i s c h a p t e r i s t o examine the d r y weight  and d i s t r i b u t i o n o f t h e v a r i o u s components o f 1 0 0 - y e a r - o l d l o d g e p o l e p i n e t r e e s , t o 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 d r y w e i g h t s characteristics, old  o f the components and s e v e r a l e a s i l y measured  tree  and t o compare the t o t a l s t a n d i n g crop o f 100-year-  l o d g e p o l e p i n e stands growing under d i f f e r e n t s i t e and s t a n d  density conditions i n Alberta.  5  Methods and Materials  Data collection  The data for this study were gathered from sample plots located i n each of three stands on the Kananaskis Forest Research Station (North Latitude 51°06', West Longitude 115°04') i n the SA 1 Section of the Subalpine Forest Region (Rowe, 1959). The three stands were on a well-drained, gently sloping s i t e , of medium site quality, at an elevation of about 4,600 feet.  The s o i l i s 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 l i s t s some of the most prominent lesser vegetation present i n 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 i s not known. No treatment was applied to Stand 3 and the condition of this stand i n 1938 i s unknown. When the present study was carried out most of the trees i n 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 i n 1938 before and after thinning. Stand 1 Characteristics  Living stems per acre Basal area per acre (sq.ft.) Volume per acre (cu. ft.) Mean dbh (in.)  Stand 2 Before Thinning  3,005 195.6 3,905 3.5  2,003 187.6 4,149 3.6  After Thinning 1,220 128.6 2,910 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 i n each of Stands 1 and 2, and a rectangular twentieth-acre plot was located i n Stand 3. The characteristics of the stands, based on plot measurements are presented i n Table 2.  Table 2. Characteristics of the three stands of 100year-old lodgepole pine. Characteristic  Stand 1  Stand 2 (Thinned)  Stand 3  Living stems per acre Basal area per acre (sq.ft.) Volume per acre (cu.ft.) Mean dbh (in.) Mean height (ft.)  1,020.0 227.7 6,356.0 6.5 54.7  290.0 152.0 5,107.0 9.8 66.5  4,960.0 156.5 2,594.0 2.2 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 i t s 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 i n moisture content (Figure 1), an a r i t h 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 Sy.x • 0.039  r « O.I37**n » 713.  20  30 Height  Source• Johnstone [ 1 9 7 0 a ]  H (ft)  2  40  in Tree [ F t . a b o v e  50 Ground]  60  70  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 i n 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 s o i l 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 i n this method of extraction, the losses, i n terms of their dryweight, were small. Data from 88 sample plots, ranging i n size from 1/20 to 1/2 acre, established i n essentially pure, 100-year-old (91-110 years) lodgepole pine stands (stands i n 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, s i t e , 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  10  10  No.  of Stems). Because the data used to derive the above equation were collected from a wide range of age, s i t e , and stand density conditions the equation was deemed to represent an average for the species i n 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), i n 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 i n 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 ) : 3  the dry weight of the stem,  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.  g)  Branch dry-weight (Y ):  h)  Stump plus root dry-weight (Y ):  6  7  the dry weight of living branches. 8  the dry weight of a 1.0-  foot stump plus roots. i)  Root dry-weight (Y ) : the dry weight of that portion 9  of the tree below ground l e v e l . 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 f a c i l i t a t e the application of a linear model and to permit comparisons with similar work reported i n the literature which most frequently use this transformation. In the following results the standard error of estimate i s expressed both in absolute units and as a percentage of the mean, the latter being i n parentheses. Because the standard error of estimate  13  determined cannot  be  from  the r e s i d u a l mean square  of a l o g a r i t h m i c equation  transformed back to an a r i t h m e t i c s c a l e w i t h o u t b i a s ,  s t a n d a r d e r r o r s o f e s t i m a t e were c a l c u l a t e d by  where: Y = observed 5  the  the f o l l o w i n g f o r m u l a :  v a l u e of the dependent v a r i a b l e  Y = e s t i m a t e d v a l u e o f the dependent v a r i a b l e n = number of o b s e r v a t i o n s  S i m i l a r l y , an e s t i m a t e d c o e f f i c i e n t of d e t e r m i n a t i o n ( r ) was  obtained  2  by  the f o l l o w i n g f o r m u l a : R  2  2  or  r = SStotal-SSresid. SStotal  where:  S S t o t a l = sum SSresid.=  I  Because the squared  o f squares (Y-Y)  of untransformed  Y  2  d e v i a t i o n s of l o g a r i t h m i c v a l u e s r a t h e r  than a r i t h m e t i c v a l u e s a r e minimized,  the use of l o g a r i t h m i c e q u a t i o n s ,  d e r i v e d by  the l e a s t squares methods, r e s u l t s i n a s y s t e m a t i c under-  estimate.  A method of overcoming t h i s problem was  (1944)  based  suggested  by Meyer  on the f o l l o w i n g f o r m u l a : 2 C  F  =  10  1.1513-<*  where: CF = c o r r e c t i o n a  factor  = s t a n d a r d e r r o r of e s t i m a t e of a l o g a r i t h m i c e q u a t i o n i n terms of l o g a r i t h m s determined  from  (i.e.,  as  the r e s i d u a l mean square)  Figure 3 presents this correction factor  (CF) g r a p h i c a l l y .  e s t i m a t e s from a l o g a r i t h m i c e q u a t i o n need merely  Parameter  be m u l t i p l i e d  by  the a p p r o p r i a t e c o r r e c t i o n f a c t o r f o r t h a t e q u a t i o n to o b t a i n u n b i a s e d  14  estimates. As can be seen from Figure 3 the application of the correction factor becomes c r i t i c a l 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  R e s u l t s and D i s c u s s i o n Component d r y - w e i g h t s  o f 100-year-old  lodgepole pine trees  T a b l e 3 p r e s e n t s the means, s t a n d a r d d e v i a t i o n s , and minimum and maximum v a l u e s o f the independent  T a b l e 3.  Mean  Average s t a n d d e n s i t y  (Stands  2  2  2  Minimum value  Maximum value  2.1 7.5 10.3 2.0 4.9 6.6 32.7  4.0 45.0 8.0 2.5 75.0 20.1 16.0  13.4 81.7 54.0 14.7 100.0 50.8 179.6  2 ,581.0  760.0  13,772.2  1.0 6.7 4.8 1.0 2.8 5.7  0.6 5.7 1.6 0.5 3.7 0.4  6.6 41.0 27.0 6.5 18.2 43.6  205.3  2.1  1,781.6  s t a n d d e n s i t y (Stand 3) - 221 o b s e r v a t i o n s  Diameter (D) ( i n . ) 2.2 H e i g h t (H) ( f t . ) 18.8 Crown l e n g t h (CL) ( f t . ) 9.4 2.5 Crown w i d t h (CW) ( f t . ) H e i g h t to l i v e crown (HLC) ( f t . ) 9.3 5.8 D i a m e t e r squared ( D ) ( i n ) . Diameter squared times h e i g h t (D H) (in. • ft.) 141.9 2  2  2  The of  Stand dev.  1 and 2 pooled) - 85 o b s e r v a t i o n s  D i a m e t e r (D) ( i n . ) 7.1 60.4 H e i g h t (H) ( f t . ) Crown l e n g t h (CL) ( f t . ) 21.6 Crown w i d t h (CW) ( f t . ) 5.4 Stump age (AGE) ( y r s . ) 92.4 H e i g h t to l i v e crown (HLC) ( f t . ) 3 8 . 8 Diameter squared ( D ) ( i n ) . 54.3 Diameter squared times h e i g h t (D H) (in. • ft.) 3,498.4 High  i n the a n a l y s e s .  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 o f the independent v a r i a b l e s f o r 100-year-old lodgepole pine t r e e s .  Independent variables  2  v a r i a b l e s used  2  means, s t a n d a r d d e v i a t i o n s , and maximum and minimum  t h e dependent v a r i a b l e s a r e p r e s e n t e d i n T a b l e 4.  values  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 o f the dependent v a r i a b l e s for 100-year-old lodgepole pine t r e e s .  Independent variable  Average  Mean  stand density  Minimum value  Maximum value  (Stands 1 and 2 p o o l e d ) - 85 o b s e r v a t i o n s  T o t a l t r e e dry-weight T o t a l above-ground dry-weight T o t a l stem dry-weight M e r c h a n t a b l e stem dry-weight Stem b a r k d r y - w e i g h t Needle d r y - w e i g h t Branch d r y - w e i g h t Stump p l u s r o o t dry-weight Root d r y - w e i g h t High stand d e n s i t y  Stand dev.  (Stand  T o t a l t r e e dry-weight T o t a l above-ground dry-weight T o t a l stem dry-weight Needle d r y - w e i g h t Branch d r y - w e i g h t Stump p l u s r o o t dry-weight  ( l b ,.) 372.7 (Y2) ( l b .) 311.4 ( Y 3 ) ( l b .) 268.9 ( Y 4 ) ( l b .) 237.3 <*5> ( l b • ) 31.2 ( Y 5 ) ( l b • ) 14.0 ( Y 7 ) ( l b .) 21.8 ( Y 8 ) ( l b .) 57.4 ( Y 9 ) ( l b .) 51.5  (Yi)  280 .1 226 .2 179 .8 186 .4 29 .7 11 .7 30.8 49 .8 48 .1  92.3 64.3 58.9 0.0 5.3 1.0 0.4 13.2 9.1  1,530.3 1,238.1 944.3 930.0 170.8 61.2 194.9 292.2 283.5  3) - 221 o b s e r v a t i o n s (Yi) (lb (Y2) (lb (Y3) (lb (Y6) ( l b (Y7) (lb (Y8) (lb  .) 20.8 • ) 16.8 .) 12.5 1.4 .) 2.9 .) .) 4.0  27 .2 22 .2 16 .0 2.2 4 .3 5.1  0.8 0.6 0.4 0.02 0.04 0.2  214.8 173.8 125.4 16.1 32.3 41.0  72 o b s e r v a t i o n s were used f o r r o o t p l u s stump w e i g h t , r o o t w e i g h t , and t o t a l t r e e w e i g h t , t h e r e f o r e , the dependent v a r i a b l e s a r e n o t additive.  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 u n t r a n s f o r m e d  dependent and independent v a r i a b l e s a r e p r e s e n t e d i n T a b l e 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 variables 1  Dependent variables Y2  Y«+  Y3  Y5  Y6  Y7  Y8  Y9  0.814 0.713 0.694 0.867 0.034 -0.251 0.881 0.902  0.914 0.825 0.753 0.885 0.060 -0.251 0.960 0.972  0.899 0.809 0.744 0.879 0.055 -0.256 0.950 0.965  0.873 0.812 0.832 0.709 0.513 0.936 0.943  0.897 0.847 0.841 0.697 0.581 0.959 0.970  3 Average stand density (Stands 1 and 2 pooled) - 85 observations  D H CL  cw  AGE HLC D2 D2H  0.966 0.894 0.796 0.888 0.155 -0.222 0.989 0.992  0.961 0.882 0.783 0.894 0.098 -0.235 0.989 0.993  0.975 0.912 0.798 0.873 0.173 -0.204 0.989 0.989  0.979 0.917 0.800 0.879 0.180 -0.202 0.990 0.989  . 0.870 0.852 0.833 0.831 0.074 -0.328 0.900 0.922  0.921 0.841 0.782 0.883 0.185 -0.262 0.940 0.940  High stand density (Stand 3) - 221 observations D H CL CW HLC D2 D2H p.05 1 2 3  0.922 0.871 0.859 0.725 0.606 0.983 0.992 signif. level  0.921 0.871 0.858 0.727 0.608 0.982 0.990 ^  y Q  *  « 0.232  0.924 0.879 0.853 0.721 0.656 0.983 0.992 r  0.873 0.812 0.832 0.709 0.513 0.936 0.943  83  = 0.216  9  r  - 0.137 y  See Table 3 for definition of abbreviations. See Table 4 for definition of abbreviations. 72 observations were used for root plus stump, root, and total tree dry-weights.  19  As can be seen by the results presented i n 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 independent 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 i n 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. Therefore i t i s 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 i n Table 6 this varied from less  Table 6. Dry-weight allometric relationships for 100-year-old lodgepole pine trees.  Dependent variable ( l b . ) 1  Intercept  Regres. coeff.  Independent var.(in. ft.)  r2  Sy.x (lb.)  CF  AD  72 85 85 85 85 85 85 72 72  0.987 0.985 0.980 0.979 0.849 0.875 0.860 0.936 0.923  32.0 28.1 25.8 27.4 11.6 4.2 11.6 12.7 13.4  1.003 1.004 1.005 1.000 1.053 1.057 1.145 1.013 1.017  -0.002 -0.005 -0.000 0.000 -0.054 -0.027 -0.104 -0.024 -0.030  221 221 221 221 221 221  0.981 0.984 0.988 0.876 0.876 0.908  3.8 2.8 1.7 0.8 1.5 1.6  1.025 1.021 1.015 1.236 1.236 1.053  -0.038 -0.021 -0.013 -0.151 -0.150 -0.085  No. observ.  Average stand density (Stands 1 and 2 pooled) Logi o Yi Logi o Y2 Logi o Y3 YLogi o Ys Logi o Y6 Logi o Y7 Logio Y8 Logic. Y9  -0.966 -0.998 -0.889 -12.626 -2.584 -2.951 -4.126 -1.880 -2.104  0.997 0.985 0.938 0.071 1.140 1.148 1.509 1.022 1.070  2  Logio D H 2 Logi o D H 2 Logio D H D2H 2 Logi o D H 2 Logi o D H 2 Logi o D H Logi o D2H 2 Logi o D H  High stand density (Stand 3) Logi o Yi Logi o Y2 Logi o Y3 Logi o Y6 Logi o Y7 Logio Y8  -0.578 -0.749 -0.823 -2.108 -1.806 -1.123  0.887 0.923 0.901 1.023 1.023 0.806  2 Logi o D H Logi o D2H 2 Logi o D H 2 Logi o D H 2 Logi o D H 2 Logi o D H  See Table 4 for definition of abbreviations.  21  F i g u r e 4. T O T A L T R E E •AVERAGE  WEIGHT - O H  STAND  LODGEPOLE  DENSITY  ALLOMETRY  Figure 3 . TOTAL  OF  D2H  100-YEAR-OLD  1  ABOVE-GROUND  ALLOMETRY  100-YEAR-OlD  PINE.  COMPONENT  O F 'AVERAGE  LODGEPOLE  STAND  WEIGHTDENSITY1  PINE.  i J J 6  U9 Y >0.«97 | 0  )  • 0.9W t o j ^ o V ( » 3 «.l-< 99«  I «, O H in lt.1-0.966 0  5  3  »• 7J  r'"0.9aj «"»S  S r ,-330 lb. l i t *>>  v.*  ».l lb.  (9.0 %\  0Vl (in. -HI 1  IT* (KI/11.1  F i g u r e 6. S T E M  WEIGHT- D H  'AVERAGE OLD  STAND  LODGEPOLE  ALLOMETRY  DENSITY*  OF  F i g u r e 7. M E R C H A N T A B L E  100" Y E A R -  PINE.  STEM  WEIGHT - D^H  RELATIONSHIP  OF'AVERAGE  100-YEAR-OtD  LODGEPOLE  STAND  DENSITY'  PINE.  /  £  1 ^ 1 0 J " 0»3» l o g p ^ H l i , Y  » » 0.980 3  3  too  VA- 0 . 0 7 1 D H [ i n * l l . ) - 1 J . 6 2 6 3  ll.) - o.»«»  r " 0.979  « • »5  3  >'•• ">•« 10  o3*  n  « as  S,..'"' lb.  a»o  woo  6ooo O'H t i - 3 i . . ]  eooo  uooo  12.000  uoco  22  Figure 8. S T E M B A R K W E I G H T - D ^ H  ALIOMETRY  OF  Figure 9.  NEEDLE "AVERAGE  LODGEPOLE  PINE.  OLD  WEIGHT-0 STAND  LODGEPOLE  H  ALIOMETRY  DENSITY' PINE.  OF  100 - Y E A R -  23  ngw* 12. tOOT WEIGHT - D H  ALLOMETRY Of  'AVERAGE STAND DENSITY' 100-YEAROLD lOOGEFOLE PINE.  24  Figure 13. TOTAL TREE WEIGHT-D H ALLOMETRY OF 'HIGH STAND DENSITY' 100"YEAR-OLD LODGE POLE FINE. 2  F i g u r e 14. T O T A L D  2  H  ABOVE - GROUND ALLOMETRY  WO-YEAR-OLD  OF  COMPONENT  "HIGH  STAND  LODGEPOLE  PINE.  WEIGHT-  DENSITY'  4k *4F^  2  4k  I|  4k  3EK,  *  I  A  1  A  oi J  Jr  A  f  41  * 4k A >  <  u>QioV * 7 Log, ,0 M[iii. -#t.J-Q 578 < -pi  *  4  A  «r  2  ft  / .  a  4k  0.749  IOSXJ'J" " 0  4k  »'0.984 n14.7%; 5 . - 2.8ft.( 3  $ . - 3.8 r  lb.  18.3% J  R  0 M [m. 5  Figure  15. S T E M  WEIGHT-D  STAND POLE  DENSITY'  llj  J  2  H  ALLOMETRY  100-YEAR-OLD  OF  "HIGH  Figure  16.  NEEDLE *HIGH  LODGE-  WEIGHT - D  STAND  LO0GEPOLE  PINE.  2  H  ALLOMETRY  DENSITY'  OF  100-YEAR - OLD  PINE.  023 to.DH - 2.108 J  ln  i> -231  .'-0.876 S . „•<>.« Y  o. [5i<•*>]  A / k  A  » K i  4k 4k  !  "iv. 7-  *  i  t  *r •  4k 4k . A Ay A A V A * *m A / % a * • A  A  4  AAA  4k 4k  4k  jt  4k  U e „ T , «0  .'-0.988  AygOkA  4  A  A  A  A  i AAA/A  4b  n "22  A  A\  A  k n  A  A  A\ A  A * / * A W f e A - A  •01 log ^O'M - 0 823  0'H tm?- H . J  /4».  •*  1  Z  r  A  A  25  f i g u r e 17.  BRANCH 'HIGH  WEIGHT - D  STAND  IODCEPOLE  Ij'  ,'.0 v . -  H  ALLOMETRY  OF  100 - Y E A R -  Figure  18. S T U M P OF  O l D  PINE.  PLUS  'HIGH  ROOT  STAND  LODGEPOLE  W E I G H T - D DENSITY'  2  H  100  ALLOMETRY " Y E A R - O L D  PINE.  TOOTLOO^D'H - I.IVI  1.0 23 l o s ^ D ^ M -1 B06  •76  1.3  2  DENSITY'  r'- 0.901  „ -111  b. [40.  lb. [ 5 1 . 7 % ] *  ft A *  A  i  L  ft * *  *  *  A A  i  A  A,  *  '  *  * A/  A  i  A1*  •  A A  *AT  «k 4ft A  0H J  [in. llj 3  A  .  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, Y 2 , and Y 3 )  this  underestimate may be ignored. In other instances, particularly branch dry-weight ( Y 7 ) , this systematic bias i s 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 i n an over-estimate of approximately 8.5 percent for needle and branch dry-weights. This may be due to the high variability i n 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 i t s 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 i n the plot were  measured and are included i n 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 i n Appendix I I .  27  Table 7. Dry weight of tree components and total standing crop for three stands of 100year-old lodgepole pine. Component  STAND 1 STAND 2 (oven-dry pounds per acre)  Needles 11,619 Branches 15,209 Stem 193,045 Total above l'-stump 222,613 Roots plus stump 39,203 Total a l l components 262,941 1  (4.4%) (5.8%) (73.4%) (84.7%) (14.9%)  13,324 22,589 136,134 173,267 31,585 206,090  STAND 3  (6.5%) 7,688 (11.0%) 15,398 (66.1%) 61,816 (84.1%) 83,207 (15.3%) 19,228 101.637  (7.6%) (15.1%) (60.8%) (81.9%) (18.9%)  The figures are not additive because they were determined from independent least-square logarithmic regression equations.  As can be seen from the results presented i n the preceding table (Table 7) there is a marked difference i n the distribution of dry-weight by components i n the three sample stands.  In Stand 2, which  had been thinned previously, a greater proportion of the total dryweight was contained i n the foliage and branches than i n i t s unthinned counterpart (Stand 1).  This suggests that the thinning resulted i n  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 i n Stands 1 and 2 (11.39 and 45.94 pounds per tree, respectively), the greatest proportion of crown materials per acre (Table 7) was supported by the highly suppressed trees i n Stand 3. Because the high stand density probably resulted i n 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 l i g h t , l i t t l e energy was a v a i l able for wood production.  The relationships between photosynthesis  28  and  l i g h t i n t e n s i t y a r e d i s c u s s e d by Kramer and K o z l o w s k i  Nichiporovich  (1967) , Wassink (1968) , and Monsi  Component dry-weights  of 100-year-old  (1960) ,  (1968).  lodgepole pine  stands  T a b l e 8 p r e s e n t s the s t a n d c h a r a c t e r i s t i c s used v a r i a b l e s i n the a n a l y s i s o f e s t i m a t e d l o d g e p o l e p i n e stands  T a b l e 8.  from  independent  t o t a l s t a n d i n g crop o f 1 0 0 - y e a r - o l d  Alberta.  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 o f the independent v a r i a b l e s from e i g h t y - e i g h t stands o f 100-yearold lodgepole pine.  Mean  Independent variable  Age (AGE) ( y r . ) 98.0 AI (AI) (%) 100.0 Kl (Kl) ( f t . ) 56.8 Number o f stems/acre (N) 884.1 Average diameter (D) ( i n . ) 6.0 B a s a l a r e a / a c r e (BA) (s_q.ft.) 162.9 Average s t a n d h e i g h t (H) ( f t . ) 52.2 B a s a l a r e a times ave. h e i g h t 8,531.4 (BA-H) ( f t . - f t . ) B a s a l a r e a d i v i d e d by a v e . h e i g h t (BA/H) ( f t . / f t . ) 3.3 Ave. diameter d i v i d e d by a v e . h e i g h t (D/H) ( i n . / f t . ) 0.11 2  2  The  as  estimated  Standard dev.  Minimum value  Maximum value  6.0 9.2 8.2 402.1 1.4 28.2 11.3  91.0 80.0 44.0 260.0 3.6 108.6 24.1  110.0 126.0 81.0 2,140.0 9.5 222.8 72.9  2,433.5  3,198.1  14,347.7  0.9  1.8  5.5  0.02  0.09  0.15  component dry-weights  p e r a c r e used as the  dependent v a r i a b l e s i n t h e a n a l y s i s o f t o t a l s t a n d i n g crop o f 100-yearold  l o d g e p o l e p i n e stands from A l b e r t a a r e p r e s e n t e d i n T a b l e 9.  29  Table 9. Statistical characteristics of the dependent variables from eighty-eight stands of 100year-old lodgepole pine.  Dependent variable (lb./ac.) Needle dry-weight Branch dry-weight Stem dry-weight Above-ground dry-weight Root plus stump dry-weight Total tree dry-weight  (Yio) (Yu) (Y12) (Y13)  (YiO  (Y15)  Mean  Standard dev.  Minimum value  Maximum value  6,612 9,154 135,163 152,621 27,203 180,590  1,974 4,189 30,977 36,935 6,893 44,349  3,622 3,719 82,969 93,643 16,279 109,914  11,266 19,894 214,073 246,076 44,454 292,460  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 variables 1 AGE AI Kl N D BA H BA-H BA/H D/H  Dependent variables Y13 Ym  Yio  Y11  Y12  0.262 0.407 0.595 -0.463 0.762 0.601 0.754 0.949 -0.277 0.053  0.314 0.238 0.656 -0.643 0.872 0.372 0.756 0.816 -0.366 0.257  0.175 0.556 0.444 -0.170 0.537 0.796 0.642 0.976 -0.117 -0.176  0.201 0.521 0.492 -0.255 0.606 0.751 0.683 0.981 -0.164 -0.116  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.  0.218 0.494 0.523 -0.313 0.652 0.716 0.707 0.979 -0.197 -0.072  Yis 0.207 0.513 0.502 -0.275 0.622 0.740 0.691 0.981 -0.175 -0.101  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 ( Y i i ) , are most closely associated with the combined variable of stand basal area times the average height of the trees i n 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 i n a negative correlation (r = -0.315) between number of stems and the combined variable (BA'H). It i s 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 i n 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 F t • ACM  titm 20.  AND STAND 1ASAL AREA TIMES MEAN STAND HEIGHT FO*  THE  RELATIONSHIP BETWEEN ESTIMATED DRY IRANCH WEIGHT FER  ACRE AND STAND IASAL AREA TIMES MEAN STAND HEIGHT FOR  STANDS OF KX>-TEA«-OtO IODGEFOIE FINE.  STANDS OF 100-YEAR-OLD IODGEFOIE  FINE.  t  1 I  i  a.ooo  IMOO  r^ura  at.  13.000  • A A (It'-tt. )  %» H (h.'-ti.)  THE RELATIONSHIP MTWEEN ESTIMATED DRY STEM WEIGHT PER Figur»22. THE RELATIONSHIP BETWEEN ESTIMATED DRY ABOVE "GROUND WEIGHT PER ACRE AND STAND BASAL AREA TIMES MEAN STAND HEIGHT r*OR ACRE AMD STAND BASAL AREA TIMES MEAN STAND HEIGHT STANDS OF KW-YEAR-OLD LODGEPOLE PINE. FOR STANDS Of 100 - YEAR - OLD LODGEPOLE PINE. 130.000  v - it.mi + n.43 •* ft n  .'•MM  tow  r - l i . j j o » t u t t i * fi )3  m-tt  upoo •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"YEAROLD LODGEPOLE PINE. 300.000-  260.000.  i 220.000.  I  A  A  A  I  O  A  180.000  A  A  *  1  140.000 -  A A A  A  A  A A A A  100.000 . 200  600  ljOOO Number  1.400 of  Stems  1.800  2.200  34  heterogeneity of variance displayed i n 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 i n Table 11. The regression statistics presented are: regression coefficients (B_^) and their significance (F . ) , number of observations (No.).coefficients of determinations xi (R 2 ), and standard errors of estimate (Sy.x). The results presented i n 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 w i l l contain a greater total standing crop than a stand of a large number of small trees when the basal area of the two stands i s equal. components.  .  These relationships were characteristic for a l l  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 variable (lb./ac. )  Independent variables and their significance (Xi = N, X2 = BA, X3 = A I or Kl) Regression coefficients and F-ratios 80  FX!  81  Fx2  82  No.  83  R  Sy.x (lb.)  88 88 88 88  0.945 0.867 0.603 0.413  10,596 16,393 28,266 34,383  88 88 88 88  0.960 0.911 0.867 0.259  9,060 13,351 16,393 38,635  Fx3  X3 = AI Yis Y15 Y15 Y15  -120,237 -3,337 -107,221 -68,396  -68.9 -66.4  520.8 203.1  -43.6  21.7  1,308.6 1,489.0 1,000.5  811.0 503.7 72.9  1,485 .1  119.4  1,247 .8 2,875 .4  11.9 48.8  1,416.1 1,286.3 1,489.0  1,461.8 632.6 503.7  2,228 .0 3,292 .4  194.3 349.4  3,091 .6  21.0  X3 = Kl Y15  Yis Yis Y15  -146,179 -216,015 -3,337 -5,165  -34.5  100.6  -66.4 11.5  203.1 0.7  p. 01 s ignificance level Fj_  -=  8  6.95  ^1 t 8 5  =  • 6.95  36  Conclusions  It i s 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 i s probable that these estimates would be improved had the  quantity of bark and branch components been obtained by direct measurement. Systematic underestimates w i l l 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 i s greatest for the crown components. Meyer's (1944) method for overcoming this problem i s generally satisfactory but may, as i n the case of the crown components, result i n 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 i s influenced by extremes i n stand density. This i s particularly 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 i n terms of conventional height-age relationships may  37  r e s u l t i n erroneous  conclusions.  However, from t h e r e s u l t s  presented  i n T a b l e 7, comparisons o f p r o d u c t i v i t y i n terms o f t o t a l t r e e biomass are also unadvisable.  C o r r e c t i o n f o r the o b v i o u s o m i s s i o n o f the biomass  c o n t r i b u t e d by l e s s e r ground v e g e t a t i o n would n o t improve the e s t i m a t e s i g n i f i c a n t l y because i n a l l t h r e e s t a n d s t h i s v e g e t a t i o n was n e g l i g i b l e . These r e s u l t s suggest stand dry-matter or  t h e r e f o r e , t h a t one must be c a u t i o u s i n comparing  p r o d u c t i o n when extremes i n s t a n d d e n s i t y a r e encountered  when a major s t a n d d i s t u r b a n c e has o c c u r r e d a f t e r the age a t which  the s t a n d c a n f u l l y compensate f o r t h a t d i s t u r b a n c e . Although old  t h e biomass e s t i m a t e s o f the 88 stands o f 100-year-  lodgepole pine represent, a t b e s t , a f i r s t approximation  the a n a l y s i s does suggest  (Table 9 ) ,  t h a t , w i t h the p o s s i b l e e x c e p t i o n o f b r a n c h  biomass p e r a c r e , r e l i a b l e e s t i m a t e s o f component biomass can be o b t a i n e d u s i n g the combined v a r i a b l e o f b a s a l a r e a p e r a c r e times mean s t a n d height  ( T a b l e 1 0 ) . The s t r o n g i n v e r s e c o r r e l a t i o n between b r a n c h  biomass  and numbers o f stems p e r a c r e and the s t r o n g p o s t i v e c o r r e l a t i o n o f b r a n c h biomass and average nature o f the s p e c i e s . is  stand diameter  intolerant  The r e s u l t s i n d i c a t e t h a t the amount o f f o l i a g e  c l o s e l y a s s o c i a t e d w i t h d r y - m a t t e r p r o d u c t i o n ( T a b l e 10) thus  porting Stiell  the r e s u l t s o f M a r : M o l l e r (1966).  Unlike MarrMoller  f o l i a g e biomass was n o t apparent, in  r e f l e c t s the r e l a t i v e  (1947), B a s k e r v i l l e (1947) and S t i e l l  Mar:Moller  (1965a), and  (1966), a c o n s t a n t  a l t h o u g h the range i n d e n s i t y examined  t h i s study was r e l a t i v e l y narrow.  crop was o b s e r v e d .  sup-  S i m i l a r l y , no maximum t o t a l s t a n d i n g  (1947) h y p o t h e s i z e d  that a f t e r f u l l  site  occupancy h a d been a c h i e v e d growth becomes s t a t i c and changes i n s t a n d d e n s i t y o n l y a l t e r the d i s t r i b u t i o n and n o t the amount o f growth.  The  r e s u l t s p r e s e n t e d i n T a b l e 11 tend t o r e f u t e t h i s h y p o t h e s i s f o r l o d g e p o l e pine.  The r e s u l t s p r e s e n t e d  t h e r e i n demonstrate t h a t , on a g i v e n  site  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 p r e c e d i n g c h a p t e r o f t h i s  c o n s i d e r a b l e a t t e n t i o n has been devoted p r o d u c t i v i t y , i n terms o f d r y - m a t t e r systems.  Surprisingly, relatively  thesis  t o s t u d i e s o f the t o t a l  p r o d u c t i o n , o f many f o r e s t  eco-  few s t u d i e s ( O v i n g t o n , 1957; T a d a k i  e t a l . , 1962; B a s k e r v i l l e , 1965a, 1966; Ando, 1965; K i r a and S h i d e i , 1967;  B e l l a and J a r v i s , 1967; Satoo,  e t a l . , 1970; J o h n s t o n e ,  1967; Osborn, 1968; P e t e r s o n  1971; O l s o n , 1971; S t e i n b e c k and May, 1971;  M o i r and F r a n c i s , 1972; P o l l a r d , 1972; H e g y i , 1972) have been to comparing the d r y - m a t t e r  production of a s i n g l e species over a  range o f s i t e and/or s t a n d c o n d i t i o n s . s t u d i e s which compare the d r y - m a t t e r grown on s i m i l a r s i t e s P o s t , 1970).  devoted  Fewer y e t a r e the number o f  production of d i f f e r e n t species  ( M a r : M o l l e r , 1947; O v i n g t o n ,  1956; Assmann, 1961;  The o b j e c t i v e o f t h i s study was t o compare, f o r two  d i f f e r e n t a r e a s , the above-ground t o t a l s t a n d i n g crop o f a f u l l y s t o c k e d s t a n d o f young l o d g e p o l e p i n e w i t h a f u l l y - s t o c k e d s t a n d o f Populus climatic  s p e c i e s o f a s i m i l a r age and grown under s i m i l a r e d a p h i c and factors.  40  Methods and M a t e r i a l s  The square  d a t a f o r t h i s study were gathered  from f o u r , one hundred  metre ( a p p r o x i m a t e l y f o u r t i e t h - a c r e ) sample p l o t s l o c a t e d i n two  areas n e a r Fox Creek, A l b e r t a i n the B o r e a l b i o c l i m a t i c zone 1959).  In A r e a 1 (North L a t i t u d e 54°14', West L o n g i t u d e  paired lodgepole pine  (Rowe,  116°17*),  ( P l o t 4) and t r e m b l i n g aspen (Populus  tremuloides  Michx.) ( P l o t 5) p l o t s were e s t a b l i s h e d on an upper s l o p e o f 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 ,  the dominant u n d e r s t o r y v e g e t a t i o n was Viburnum-Linneae. of  white  s p r u c e and w i l l o w  and  A s m a l l number  ( S a l i x L.) t r e e s were a l s o p r e s e n t .  A l l pine  and aspen t r e e s were about 23 y e a r s o l d . In A r e a 2 (North L a t i t u d e 54°17', West L o n g i t u d e paired  lodgepole pine  116°10'),  ( P l o t 6) and balsam p o p l a r (Populus b a l s a m i f e r a L )  ( P l o t 7) p l o t s were e s t a b l i s h e d on a lower s l o p e a t an e l e v a t i o n o f about 2,265 f e e t .  The s o i l  i s a g l e y e d grey wooded over l a c u s t r i n e c l a y , and  the dominant u n d e r s t o r y v e g e t a t i o n was L o n i c e r a - H e r a c l e u m . of  white  s p r u c e , w i l l o w and w h i t e b i r c h  were a l s o p r e s e n t . The  A s m a l l number  ( B e t u l a p a p y r i f e r a Marsh.) t r e e s  The p i n e and p o p l a r t r e e s were about 25 y e a r s o l d .  c h a r a c t e r i s t i c s o f t h e s t a n d s , based  are p r e s e n t e d i n T a b l e 12.  on p l o t measurements,  41  Table 12.  Characteristics of the four lodgepole pine and Populus stands.  Characteristic  Plot 4 Plot 5 Plot 6 (l.pine) (t.aspen) (l.pine)  Living stems per acre Basal area per acre (sq.ft.) Mean dbh (in.) Range of dbh (in.) Mean height (ft.)  6,515 135.7 1.9 0.4-3.8 19.1  7,972 3,683 121.3 142.0 1.5 2.5 0.3-3.8 0.7-4.8 20.2 28.6  Plot 7 (b.poplar) 2,671 169.9 3.0 0.2-6.0 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 i n each area, from which the following 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  f.  Age  g.  Stump fresh-weight  h.  Stem fresh-weight  i.  Fresh, and dry weights of branches greater than 2 cm.  (CW)  (AGE)  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 l e v e l , 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 dryweight allometric relationships were obtained using methods similar to those outlined i n 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 following 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 freshweight 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 i n 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 i n 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 l e v e l . 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 (Y 3 ): 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 ) :  f)  Dry dead branch weight (Ye): the dry weight of a l l dead  the dry weight of a l l green needles.  44  branches. g)  Above-ground living dry-weight ( Y 7 ) : the dry weight of a l 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 (Y 9 ): 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 + Y 9 ) .  Results and Discussion  Table 13 presents the means, standard deviations, and minimum and maximum values of the independent variables used i n the regression analyses.  45  Table 13. Statistical characteristics of the independent variables from twenty young lodgepole pine trees.  Independent variable  Mean  Diameter 2.1 (D) (in.) Height (H) (ft.) 25.4 Crown length (CL) (ft.) 11.1 Crown width (CW) (ft.) 2.5 Age (AGE) (yrs.) 25.6 Diameter at crown base (DCB) (in.) 1.6 Height to live crown (Ht.LC) (ft.) 14.3 Diameter squared (D ) (in. 2 ) 4.9 Diameter squared times height (D2H) ( i n . 2 f t .) 139.1  Standard deviation 0.8 5.2 3.5 1.1 2.8 0.7 2.8 3.3 112.6  Minimum value  Maximum value  1.0 17.5 5.5 1.0 19.0 0.6 9.5 1.0  3.4 33.6 16.9 5.4 29.0 3.0 18.6 11.6  17.5  388.4  Dry-weight relationships  The means, standard deviations, and maximum and minimum values of the dry-weight dependent variables are presented i n Table 14.  Table 14. Statistical characteristics of the dry-weight dependent variables from twenty young lodgepole pine trees.  Dependent variable Dry stump wt. Dry bole wt. Dry stem wt. Dry branch wt. Dry needle wt. Dry dead branch wt. Above-ground living dry-wt.  (Yi) (Y2) (Y3) (YO (Ys) (Ye) (Y7)  (lb.) (lb.) (lb.) (lb.) (lb.) (lb.) (lb.)  Mean  Stand. Min. Max. dev. value value  0.9 10.7 11.7 0.9 1.2 0.5 13.7  0.6 7.9 8.4 0.8 1.1 0.3 10.2  0.2 1.4 1.6 0.1 0.1 0.1 1.8  2.0 29.8 31.9 3.0 4.0 0.9 38.9  46  Simple correlation coefficients (r) between the independent and dependent variables are shown i n Table 15. Of the independent variables analyzed, tree diameter squared (D2) was most closely associated with four of the dependent variables (above-ground living dryweight, 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 characteristics 1 D H CL CW AGE DCB Ht.LC D2 D2H  Yj  Y2  0.975 0.791 0.839 0.856 0.627 0.975 0.424 0.973 0.935  0.961 0.854 0.875 0.857 0.562 0.974 0.499 0.983 0.980  Component dry-weights Y3 Y,, Y5 0.964 0.852 0.874 0.859 0.568 0.976 0.495 0.985 0.980  0.915 0.831 0.846 0.826 0.494 0.940 0.491 0.959 0.975  0.930 0.772 0.794 0.865 0.542 0.948 0.445 0.960 0.948  Y6  Y7  0.829 0.836 0.841 0.767 0.575 0.835 0.507 0.781 0.762  0.961 0.846 0.868 0.861 0.562 0.975 0.492 0.985 0.980  . 05 signif. level r 18= 0.444 See Tables 13 and 14 for description of abbreviations. 1  Because of the high correlations among the independent variables, l i t t l e improvement i n prediction r e l i a b i l i t y 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  t r e e above-ground i n the sample p l o t s  the f o l l o w i n g  r e g r e s s i o n model was s e l e c t e d : 30+  Y = Although  SiD + 3aD  the independent  significantly  2  variable  t r e e diameter  (D) o n l y c o n t r i b u t e d  t o the e q u a t i o n f o r d r y branch weight  r e t a i n e d i n the e q u a t i o n s above-ground l i v i n g  f o r needle dry-weight,  dry-weight  t o ensure  ( T a b l e 16) i t was  stem d r y - w e i g h t , and  the a d d i t i v i t y o f the e q u a t i o n s .  T a b l e 16 and F i g u r e s 26 to 29 show the r e g r e s s i o n e q u a t i o n s used t o e s t i m a t e t h e l o d g e p o l e p i n e component dry-weights  T a b l e 16.  R e g r e s s i o n e q u a t i o n s , and r e g r e s s i o n 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 ( F x . ) , d e r i v e d from twenty young l o d g e p o l e p i n e t r e e s used to e s t i m a t e sample p l o t l i v i n g component d r y - w e i g h t s .  Dependent variable  (lb.)  1  3  S  6  Regression c o e f f i c i e n t s and F - r a t i o s Fxi Bi  Bo 1.074 0.669 0.322 2.065  Y Yu Y Y  o f the sample p l o t s .  -1.922 -0.999 -0.669 -3.590  0.56 8.81 1.73 1.36  2.962 0.464 0.460 3.886  R Fx  (lb.)  2  24.37 35.04 15.04 29.40  1.51  0.971 0.947 0.928 0.972  p.05 significance level F j = 4.45 p.01 s i g n i f i c a n c e l e v e l See T a b l e 14 f o r d e s c r i p t i o n o f a b b r e v i a t i o n s . 17  Sy.x  2  F  :  l7  0.20 0.30 1.80  =  8.40  1  Fresh-weight  relationships  The of  means, s t a n d a r d d e v i a t i o n s , and minimum and maximum v a l u e s  the f r e s h - w e i g h t dependent v a r i a b l e s a r e p r e s e n t e d i n T a b l e 17.  48  Figure 2 6 . THE  RELATIONSHIP  WEIGHT  AND  LODGEPOLE  TREE PINE  BETWEEN  STEM  DIAMETER  DRY-  OF  Figure  27. THE  RELATIONSHIP  WEIGHT  YOUNG  AND  TREE  LODGEPOLE  TREES.  BETWEEN  NEEDLE  AND  TREE  DIAMETER  OF  PINE  2 9 . THE  RELATIONSHIP  BETWEEN  DRY-WEIGHT  AND  ABOVETREE  YOUNG OF  YOUNG  LODGEPOLE  TREES. PINE  7-  TREES.  DRY-  DIAMETER LODGEPOLE  YOUNG  4  GROUND WEIGHT  OF  DRY-  Y-0.449 - 0.999D+0.464 0 ir,- 0.20 0.(22.2%) l*-0.947 n-20  Figure RELATIONSHIP  BRANCH  DIAMETER  PINE  S,.," V5I Ib.d2.99b) 0.971 n • 20  Figure 2 8 . THE  BETWEEN  TREES.  70  6-  60  5-  50  O (in.)  01 0  ,  1  ,  ,  ,  2 3 D (in.)  ,  4  5  6  49  T a b l e 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 o f the f r e s h - w e i g h t dependent v a r i a b l e s from twenty young l o d g e p o l e pine trees.  Dependent variable  Mean  Stand dev.  F r e s h stem weight (Ya) ( l b . ) F r e s h crown weight ( Y ) ( l b . ) Above-ground l i v i n g f r e s h - w e i g h t (Yio) ( l b . )  24.7 4.0 28.7  19.3 3.6 22.8  9  Simple  Tree characteristics  Max. value  3.3 0.3 3.6  70.0 13.9 83.9  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  dependent v a r i a b l e s  T a b l e 18.  Min. value  and  a r e p r e s e n t e d i n T a b l e 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 f r e s h - w e i g h t s and s e v e r a l t r e e c h a r a c t e r i s t i c s o f twenty young l o d g e p o l e p i n e t r e e s .  Component Y  1  Y  8  9  0.958 0.877 0.871 0.825 0.560 0.955 0.545 0.988 0.996  D H CL CW AGE DCB Ht.LC D D H 2  2  fresh--weights Yio 1  0.956 0.869 0.866 0.832 0.556 0.957 0.536 0.988 0.995  0.931 0.810 0.825 0.861 0.521 0.953 0.478 0.968 0.970  .05 signif. level i , i 0.444 See T a b l e s 13 and 17 f o r d e s c r i p t i o n o f a b b r e v i a t i o n s . r  =  8  U n l i k e the r e s u l t s f o r t h e component dry-weights p r e s e n t e d i n T a b l e 18 i n d i c a t e d associated  that  ( T a b l e 1 5 ) , the r e s u l t s  combined v a r i a b l e D H  w i t h the dependent v a r i a b l e s  2  tested.  i c a l l y p r e s e n t the b e s t s i m p l e l i n e a r r e g r e s s i o n  i s most c l o s e l y  F i g u r e 30 t o 32  graph-  e q u a t i o n s developed i n  50 Figure 30. THE RELATIONSHIP D H  OF YOUNG  2  0  50  B E T W E E N STEM L O D G E P O L E PINE  100  150  200  D H 2  OF YOUNG  Figure32. THE RELATIONSHIP D H 2  OF Y O U N G  AND  TREES.  250  D2H  Figure 31. THE RELATIONSHIP  FRESH-WEIGHT  300  350  400  450  (in.2-ft.)  BETWEEN C R O W N L O D G E P O L E PINE  FRESH-WEIGHT  AND  TREES.  BETWEEN A B O V E - G R O U N D FRESH-WEIGHT A N D L O D G E P O L E PINE  TREES.  450 D  2  H(in.  2  ft.)  51  the a n a l y s e s . sample  p l o t l i v i n g component  Above-ground stands  These e q u a t i o n s  ( F i g u r e 30 t o 32) were used t o e s t i m a t e  fresh-weights.  o r g a n i c matter p r o d u c t i o n o f p a i r e d l o d g e p o l e p i n e and Populus  o f s i m i l a r ages on s i m i l a r  sites  T a b l e 19 p r e s e n t s a comparison o f the measure f r e s h - w e i g h t s by s p e c i e s o f the p a i r e d sample  above-ground  plots.  T a b l e 20 p r e s e n t s a comparison o f t h e e s t i m a t e d dry-matter  above-ground  c o n t a i n e d by t h e l i v i n g components o f t r e e s i n t h e p a i r e d  sample  plots.  T a b l e 20.  A comparison o f e s t i m a t e d above-ground d r y weight ( l b . / a c . ) o f l i v i n g components i n p a i r e d l o d g e p o l e p i n e and Populus stands o f s i m i l a r ages on s i m i l a r sites. AREA 1 Plot 4 Plot 5 (l.pine) (t.aspen)  „ .. Components  Major  Species:  50,202 5,293 7,310  stem branch foliage Minor  AREA 2 Plot 6 Plot 7 (l.pine) (b.poplar)  27,588 4,363 2,319  69,368 7,140 9,503  53,368 5,408 2,476  Species:  667  10,953  b. poplar t . aspen 1. p i n e  ( a l l components) ( a l l components) ( a l l components)  3,881  w. s p r u c e willow  ( a l l components) ( a l l components)  1,555 1,307  3,775 709  93 2,363  62,805 7,410 70,215  34,270 28,012 62,282  86,011 6,641 92,652  w. b i r c h  963  12,575  . ( a l l components)  Major s p e c i e s s u b t o t a l Minor s p e c i e s s u b t o t a l Total a l l species  61,252 61,252  52  T a b l e 19.  A comparison o f measured f r e s h weights ( l b . / a c . ) of p a i r e d l o d g e p o l e p i n e and Populus stands o f s i m i l a r ages on s i m i l a r s i t e s .  Components  Major  AREA 1 Plot 4 Plot 5 (l.pine) (t.aspen)  AREA 2 Plot 6 Plot 7 (l.pine) (b.poplar)  Species:  living: stem b r a n c h >2 cm. diam. b r a n c h <2 cm. diam. (incl. foliage)  105,967  58,551 218  152,546  111,616 166  24,792  13,266  32,598  14,667  4,588 1,943  1,447 3,223  5,160 8,822  1,351 1,022  1,406 8,296  23,454  dead: branches on l i v i n g t r e e s standing trees Minor  Species:  living: b. p o p l a r t . aspen 1. p i n e w. spruce willow w. b i r c h  (all (all (all (all (all. (all  components) components) components) components) components) components)  3,238 2,792  2,023 24,061 7,218 1,519 5,059  200 6,900  dead: 1. p i n e t . aspen  Living:  Major s p e c i e s  total  1,365  379  130,759  72,035  185,144  125,449  species  total  146,501  128,287  199,326  126,449  a l l species  total  6,531  6,035  13,982  2,752  all Dead:  ( a l l components) ( a l l components)  53  The results presented i n Tables 19 and 20 suggest that, for lodgepole pine at least, Area 2 appears to be somewhat higher i n site quality than Area 1, although the latter supported about 77 percent more trees.  Reasons for the apparent reversal of this trend i n the case of  Populus stand a)  are not clear but may be related to the following: The predominant Populus species in Area 1 was trembling aspen and the predominant Populus species i n 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 i n Area 1, and therefore, may not have utilized the site as f u l l y .  c)  Unlike Area 2, a successional stage had been reached in Area 1 wherein the minor species present i n 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 i n 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 i n the stems and branches of pine trees compared to Populus species (Table 21).  54  T a b l e 21.  P r o p o r t i o n o f t o t a l above-ground dry-weight by components o f major s p e c i e s i n study a r e a s .  Proportion Component  May  of t o t a l  above-ground  AREA 1 Plot 4 Plot 5 (l.pine) (t.aspen)  Stem Branch Foliage Total*  l  (%)  79.9 8.4 11.6 100.0 not be a d d i t i v e due  AREA 2 Plot 6 (l.pine)  80.5 12.7 6.8 100.0  to r o u n d i n g  dry-weight Plot 7 (b.poplar)  80.7 8.3 11.0 100.0  87.1 8.8 4.0 100.0  errors.  Conclusions  T h i s study demonstrates  t h a t the p r o d u c t i v i t i e s o f d i f f e r e n t  s p e c i e s a r e not c o n s t a n t among themselves can be expected w i t h i n s p e c i e s from s i t e the f i n d i n g s o f M a r : M o l l e r Post  (1970) .  The  ability  and  that considerable f l u c t u a t i o n s  to s i t e .  (1947), O v i n g t o n  These r e s u l t s  (1956), Assman (1961),  o f c o n i f e r o u s s t a n d s to out-produce  s t a n d s , i n terms o f above-ground d r y - m a t t e r p r o d u c t i o n , i s i n T a b l e 20.  This greater a b i l i t y  support  of c o n i f e r s  to produce  and  hardwood  demonstrated  o r g a n i c matter  i s probably a t t r i b u t a b l e to: a)  The  g r e a t e r amount and b e t t e r means o f d i s p l a y i n g  s y n t h e t i c m a t e r i a l s by c o n i f e r s  ( O v i n g t o n , 1956;  photoKramer  and K o z l o w s k i , 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  p e r i o d o f time d u r i n g the y e a r K o z l o w s k i , 1960).  ( O v i n g t o n , 1956;  Kramer and  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 i n 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 i s the result of the laying down of an unevenly  distributed sheath of xylem (and, to a much lesser degree, phloem) c e l l s , 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 b r a n c h w h o r l s u p p o r t i n g d e c r e a s e s or remains c o n s t a n t increases radial  the most f o l i a g e .  throughout the b r a n c h l e s s  a g a i n n e a r the b u t t .  growth may  Under c o n d i t i o n s  then p r o g r e s s i v e l y  (crowding)  rings).  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 o f  radial  The  f o l l o w i n g i s a summary  theories.  Hartig's n u t r i t i o n a l  theory  w h e r e i n growth was  r e l a t e d to  b a l a n c e between t r a n s p i r a t i o n and  assimilation.  o f t r a n s p i r a t i o n and  of food  growth a t any  point  the q u a n t i t y  The  intensity  a v a i l a b l e f o r stem  point.  Schwendener's m e c h a n i s t i c  theory  i n which b o l e  form i s dependent  upon i n t e r n a l ( t h e weight o f the stem i t s e l f ) and  external  (wind)  f o r c e s , which cause s t r e s s e s i n the stem, which i n t u r n  stimulate  the cambium.  pro-  R a d i a l growth a t any  point  i s , therefore,  p o r t i o n a l to the magnitude o f s t r e s s a t t h a t Jaccard's belief  water c o n d u c t i v i t y theory  that  the  the c o n d u c t i v i t y o f water between the r o o t s and  the  a r e a o f the stem.  c a p a c i t y o f t h e s e r i n g s and  hormonal t h e o r y  cross-sectional  dead branches reduce the therefore, i n order  conductive  to m a i n t a i n  the c r o s s - s e c t i o n a l a r e a must  enough to b a l a n c e l o s s e s r e s u l t i n g from d y i n g The  predicated  annual r i n g s i n f l u e n c e s the  Further,  conductive capacity,  which was  point. on  crown through r e c e n t  4.  the  i n the crown i s p r o p o r t i o n a l to the amount  o f f o l i a g e above t h a t  3.  radial  almost d i s a p p e a r n e a r the base ( m i s s i n g o r d i s c o n t i n u o u s  of the major  2.  then  decreases  of s e v e r e c o m p e t i t i o n  growth, s e v e r a l t h e o r i e s have been p o s t u l a t e d .  1.  b o l e , and  In suppressed f o r e s t - g r o w n t r e e s maximum  growth o c c u r s n e a r e r the top and  towards the base.  R a d i a l growth then  the  increase  branches.  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  radial  58  growth t o the uneven d i s t r i b u t i o n o f growth s u b s t a n c e s  (which  o r i g i n a t e i n the e l o n g a t i n g buds) a l o n g t h e b o l e . 5.  The p i p e model t h e o r y was d e v e l o p e d by S h i n o z a k i e t a l . (1964). The amount o f f o l i a g e  ( p h o t o s y n t h e t i c ) e x i s t i n g above any l e v e l  i n the stem i s p r o p o r t i o n a l to the c r o s s - s e c t i o n a l area o f the stem p l u s branches  (non-photosynthetic) a t that  level.  C o n s e q u e n t l y , changes i n t h e amount o f p h o t o s y n t h e t i c organs above any g i v e n l e v e l i n t h e b o l e would be r e f l e c t e d by p r o p 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 a r e a o f the nonp h o t o s y n t h e t i c organs a t t h a t  level.  Comprehensive r e v i e w s o f these t h e o r i e s have been p r e s e n t e d by Onaka (1950, a and b ) , Young and Kramer (1952), F a r r a r (1963), S h i n o z a k i e t a l . (1964), H a l l the p h y s i o l o g i c a l p r o c e s s e s which it  (1961), L a r s o n  (1965), and Heger (1965).  c o n t r o l form a r e n o t y e t f u l l y  i s a p p a r e n t t h a t the l i v e crown i s b a s i c to a l l o f t h e s e stem  Although understood, form  theories. The purpose o f t h i s study i s to i n v e s t i g a t e some o f t h e w i t h i n and among t r e e v a r i a t i o n s r e l a t e these v a r i a t i o n s  i n t h e growth o f l o d g e p o l e p i n e t r e e s , and t o  to some crown c h a r a c t e r i s t i c s .  Methods and M a t e r i a l s  Data  collection  The d a t a were g a t h e r e d from 20 even-aged  (100-year-old) l o d g e p o l e  p i n e t r e e s growing i n Stands 1 and 2 d e s c r i b e d i n Chapter 2 o f t h i s Ten  trees  thesis.  (Trees numbered from 1 t o 10) were grown i n an u n d i s t u r b e d s t a n d  59  (Stand 1) and the r e m a i n i n g t r e e s ( T r e e s 11 to 20) were grown i n a t h i n n e d s t a n d (Stand 2 ) .  Sample t r e e s were s e l e c t e d  to cover a wide  range o f d i a m e t e r c l a s s e s . D i s c s , about one i n c h t h i c k , were sawn from t h e stem a t 1.0 f o o t above ground  (stump h e i g h t ) , a t b r e a s t h e i g h t , a t 9.0 f e e t  above  ground, and a t 8 . 0 - f o o t i n t e r v a l s t h e r e a f t e r to t h e base o f t h e l i v e crown ( l o w e s t branch s u p p o r t i n g l i v i n g n e e d l e s ) .  One-inch  t h i c k d i s c s were a l s o  sawn from t h e stem a t 2 . 0 - f o o t i n t e r v a l s from t h e top o f t h e t r e e to t h e base o f t h e l i v e crown.  The number o f d i s c s c o l l e c t e d v a r i e d  from t r e e to  t r e e depending upon t h e h e i g h t and t h e l i v e crown l e n g t h o f t h e t r e e s .  In  t o t a l 358 d i s c s were c o l l e c t e d . In on each d i s c . bearing the  the f i e l d ,  t r e e number and c o l l e c t i o n h e i g h t were r e c o r d e d  No d i s c measurements were made a t t h a t time.  The f o l i a g e -  twigs from each 2 . 0 - f o o t i n t e r v a l o f the stem were c l i p p e d  from  l a r g e r b r a n c h p a r t s , p u t i n t o s e p a r a t e b u r l a p s a c k s , and weighed.  The  sacks were p l a c e d i n a d r y i n g shed and n e e d l e d r y - w e i g h t s were o b t a i n e d by means i d e n t i c a l  to t h o s e o u t l i n e d  w e i g h t was measured f o r t h e b r a n c h In  i n Chapter 2 o f t h i s  thesis.  No t o t a l  parts.  t h e l a b o r a t o r y , t h e d i a m e t e r i n s i d e and o u t s i d e b a r k o f each  a i r - d r i e d d i s c were o b t a i n e d to t h e n e a r e s t 1/10 i n c h w i t h a d i a m e t e r tape. For  each d i s c ,  f o u r measurements ( t o t h e n e a r e s t 1/100 m i l l i m e t r e ) were  taken o f the l a s t at the  5-year r a d i a l  a m a g n i f i c a t i o n o f 125X.  growth increment u s i n g an Addo-X machine  The f i r s t measurement was taken on a l i n e o f  mean r a d i u s and t h e r e m a i n i n g measurements were taken a t 90-, 180-, and  270- degree a n g l e s to t h i s f i r s t measurement.  The study was r e s t r i c t e d t o  the  most r e c e n t l y completed  5-year growth p e r i o d because t h i s r e p r e s e n t s  the  l e n g t h o f n e e d l e r e t e n t i o n f o r t h i s s p e c i e s and because changes  i n the  60  crown c h a r a c t e r i s t i c s o f the t r e e s made t h e v a l i d i t y  p r i o r t o t h i s 5-year p e r i o d would  o f the a n a l y s e s l e s s c e r t a i n .  have  U s i n g the average o f t h e  f o u r r a d i a l measurements, t h e d i a m e t e r o f each s e c t i o n f i v e y e a r s p r i o r was d e t e r m i n e d .  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 a r e a o f each  d i s c d e t e r m i n e d from the p r e s e n t d i a m e t e r and t h e diameter 5 y e a r s b e f o r e r e p r e s e n t e d t h e 5-year growth stem. the  Section  i n c r o s s - s e c t i o n a l a r e a a t each p o i n t  volume growth w i t h i n  average o f t h e c r o s s - s e c t i o n a l  the d i s t a n c e  between t h e s e d i s c s .  volume growth  i n the  each t r e e was determined by m u l t i p l y i n g growth o f two c o n s e c u t i v e d i s c s  times  T r e e volume growth equals the sum o f t h e  from a l l o f the s e c t i o n s w i t h i n  each t r e e .  The t o t a l age o f  each d i s c was a l s o d e t e r m i n e d . 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 o u t f o r a l l r i n g s o f each d i s c c o l l e c t e d from one t r e e grown i n each o f the two s t a n d s . T r e e 11 from Stand 2 were s e l e c t e d because of  t h e s e two t r e e s  breast  T r e e 9 from Stand 1 and  the d i a m e t e r s a t b r e a s t  c o r r e s p o n d e d most c l o s e l y w i t h t h e mean d i a m e t e r s a t  h e i g h t o f the t e n t r e e s  sampled  from each r e s p e c t i v e  stand.  d a t a from each o f the two t r e e s were c o l l a t e d and summarized tree using  height  a s p e c i a l computer  program  The  from each  f o r tree r i n g analysis.  Analysis  Two s t a t i s t i c a l  t e c h n i q u e s were used to a n a l y z e w i t h i n and among  t r e e v a r i a t i o n s i n stem growth. variance,  The f i r s t  t e c h n i q u e , an a n a l y s i s o f  was used to t e s t d i f f e r e n c e s between p l o t s , between t r e e s  p l o t s , between s e c t i o n s differences  within  ( t h e s e d i f f e r e n c e s were f u r t h e r b r o k e n down i n t o  between crown-formed  wood and c l e a r b o l e - f o r m e d wood and i n t o  61  differences between sections within the bole and within the crown). F i n a l l y , 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. t h i s analysis  i t was necessary to compare section measurements taken at the same actual or the same r e l a t i v e positions i n each tree. (1.0-foot,  This was done by using  five  4.5-foot, .9.0-foot, 17.0-foot, and 25.0-foot) measurements from  the clear-bole, and f i v e (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 r a d i a l , 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 f o r this purpose. observations,  The f i r s t analysis incorporated  a l l of the  whereas the second analysis was limited to observations  taken s o l e l y within the crown and the t h i r d analysis was based on measurements from the c l e a r bole.  A fourth multiple regression analysis was used  to r e l a t e tree volume growth and volume growth per unit of f o l i a g e to 2 several tree c h a r a c t e r i s t i c s . R values were calculated as outlined on page  Results and Discussion  P r o f i l e diagrams which i l l u s t r a t e the r a d i a l growth at each sampling point, the f o l i a g e dry-weight i n the 2.0-foot i n t e r v a l above each sampling point, and the cumulative (total) f o l i a g e dry-weight above each sampling point, are presented for each tree i n Appendix I I I .  An  i r r e g u l a r pattern of f o l i a g e d i s t r i b u t i o n 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 deviation value  Tree height (H) (ft.) Dbh (D) (in.) Diam. crown base (DCB) (in.) Dry f o l i a g e weight (DFW) (lb.) 5-yr. <radial growth (cm.) 5-yr. 'cross-sectional area growth (cm. ) 5-yr. tree volume growth (TVG) (cu.ft.) Section age (yr.) Tree volume growth/unit f o l i a g e wt. (TVG/DFW) (cu.ft./lb.)  60. 2 7.2 5.1 14. 8 0.319  8.1 2.3 2.0 11.3 0.134  10. 61  Maximum value  43. 5 3.2 2.2 1.0 0.038  77. 6 12. 9 11. 0 44. 1 0.708  8.11  0.328  63. 40  0.694 49. 1  0.525 23.3  0.041 8.0  2.191 101. 0  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 s i z e s . Table 23 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 between several tree c h a r a c t e r i s t i c s of the 20 sample trees.  63  T a b l e 23.  D  DCB  0.906 1.000  0.877 0.966 1.000  H  H 1.000 D DCB DFW TVG D D H TVG/DFW  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 s e v e r a l t r e e c h a r a c t e r i s t i c s o f t h e twenty, 1 0 0 - y e a r - o l d l o d g e p o l e p i n e sample t r e e s .  DFW  0.798 0.939 0.911 1.000  TVG  0.762 0.924 0.911 0.977 1.000  2  D  0.873 0.980 0.970 0.952 0.941 1.000  2  . 05 See  As  2  DH  0.875 0.959 0.964 0.934 0.922 0.994 1.000  x  1 Q  (Table 2 3 ) ,  character-  F o l i a g e dry-weight i s most h i g h l y c o r r e l a t e d w i t h diameter a t height  squared  A s i m i l a r ranking and  table  i s a h i g h degree o f c o r r e l a t i o n between many o f the t r e e  istics. breast  -0.143 -0.133 -0.115 -0.168 0.006 -0.132 -0.139 1.000  signif. level r = 0.444 T a b l e 22 f o r d e s c r i p t i o n o f a b b r e v i a t i o n s .  can be seen from the r e s u l t s p r e s e n t e d i n t h e p r e c e d i n g  there  TVG/DFW  2  ( D ) and t h i s r e l a t i o n s h i p i s p r e s e n t e d i n F i g u r e 33. 2  o f 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  the t r e e c h a r a c t e r i s t i c s s t u d i e d o c c u r r e d  when l o g a r i t h m i c  transform-  a t i o n s were c a r r i e d o u t .  This d i f f e r s  leaved  1966; K i r a and S h i d e i , 1967; P e t e r s o n e t a l . ,  1970)  species  (Attiwill,  from the r e s u l t s r e p o r t e d  f o r broad-  w h e r e i n d r y f o l i a g e w e i g h t was most h i g h l y c o r r e l a t e d w i t h stem  d i a m e t e r a t the b a s e o f the l i v e crown.  T r e e volume growth f o r the p a s t  5-year p e r i o d i s most h i g h l y c o r r e l a t e d w i t h d r y f o l i a g e weight ( F i g u r e 3 4 ) , w h i c h as s t a t e d p r e v i o u s l y  i s highly correlated to tree s i z e .  I n an attempt  to d e t e r m i n e the r e l a t i v e e f f i c i e n c y o f volume growth, t h e r a t i o o f volume •growth t o amount o f f o l i a g e (TVG/DFW) was r e l a t e d to s e v e r a l t r e e acteristics. there  char-  The r e s u l t s p r e s e n t e d i n T a b l e 23 and F i g u r e 35 i n d i c a t e t h a t  i s a negative  e f f i c i e n c y and t r e e  b u t 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 size.  64  Figure 33. THE RELATIONSHIP BETWEEN NEEDLE DRY-WEIGHT  Figure 34. THE RELATIONSHIP BETWEEN PAST 5-YEAR VOLUME  AND TREE DIAMETER FOR THE 20.100-YEAR-OLD  GROWTH AND PRESENT NEEDLE DRY-WEIGHT OF  LODGEPOLE PINE TREES.  20. 100-YEAR-OLD LODGEPOLE PINE TREES.  . I4-.  0  2  4  6  •  12  10  0  14  1 0  i  1  1  i  10  20  30  40  OfW  Figure 35. THE RELATIONSHIP BETWEEN VOLUME GROWTH EFFICIENCY (cu. ft. / lb. DRY FOLIAGE) AND TREE DIAMETER FOR 20.100-YEAR-OLD LODGEPOLE PINE TREES.  |  o  OJ  ?  1 -  2  4  4  8  ro  ra  14  (lb.)  r  30  —  i AO  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 i s a  decrease i n 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.  Radial growth Source  Stands Trees within stands Sections Crown vs. bole Sections within position (crowns vs. bole)  Cross-sectional area growth  df MS  F  MS  1  568  0.18ns  174  9.33**  4,307,300  5.72*  18  147,900  46.36**  541  28.93**  10,113,000  13.42**  350 1,714  18.73** 91.60**  25,195,000 189,581.000  33.43** 251.55**  180  9.62**  4,646,700  6.17**  0.34ns o.oi n s  407,760 1,526,600  0.54ns ns 2.03  nS  268,100  9 1 8  Stand X section (interaction) 9 Stand X crown vs. bole 1 Stand X sections within position (crowns vs. bole) 8 Error  162  Total  199 ** *  53,802 16.87** 283,430 88.85** 25,090  7.87** ns  2,394 146  0.74 nS 0.05  F  Section volume growth  6 0.3  nS  2,675 3,190  0.84  7.0 19  F  MS  0.35  nS  0.36  753,650  Significant at the 0.01 probability l e v e l . Significant at the 0.05 probability level. Not significant. ON  67  T a b l e 25 p r e s e n t s some s i m p l e c o r r e l a t i o n c o e f f i c i e n t s between all  o f the s e c t i o n growth measurements  i n the t r e e and s e v e r a l  t r e e and  section characteristics.  T a b l e 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 a r e a , and s e c t i o n volume growth, and s e v e r a l t r e e and s e c t i o n c h a r a c t e r i s t i c s .  Characteristic  Correlation R a d i a l growth (R. Gr.)  Tree height (H) Dbh (D) Section height •(h) Section diameter (d) S e c t i o n age (A) F o l i a g e weight i n 2 f e e t above (F) Cumulative f o l i a g e weight above (CFW) Height from top (HT) Percent of t o t a l height (PH) Form (d/D)  p.05  signif.  level  The p r e c e d i n g r e s u l t s  r  1  3 5 6  =  (Table  c o e f f i c i e n t s (r) Cross-sectional a r e a growth (C.S.A.Gr.)  S e c t i o n volume growth (S.V.Gr.)  0.443 0.618 0.426 0.063 -0.343  0.562 0.701 -0.270 0.819 0.417  0.311 0.395 -0.525 0.749 0.583  0.496  0.003  -0.280  0.347 -0.240  0.884 0.529  0.678 0.679  0.321 -0.341  -0.418 0.429  -0.600 0.566  0.105  25) i n d i c a t e t h a t , i n g e n e r a l ,  growth, be i t  r a d i a l , c r o s s - s e c t i o n a l o r volume, i s most c l o s e l y a s s o c i a t e d size.  The r e s u l t s f u r t h e r i n d i c a t e the f o l l o w i n g  d e c r e a s e d f r o m the top to the base o f the t r e e . is  true  trends.  f o r c r o s s - s e c t i o n a l a r e a and s e c t i o n volume growth.  the c o r r e l a t i o n s suggest t h a t  position within  Radial  tree growth  The r e v e r s e o f t h i s  were as expected and s u p p o r t the r e s u l t s o f the a n a l y s e s o f Further,  with  trend  These r e s u l t s variance.  r a d i a l growth i s more a f u n c t i o n o f  t h e t r e e than c r o s s - s e c t i o n a l  a r e a growth, which i s most  68  c l o s e l y a s s o c i a t e d w i t h the s i z e o f t h e t r e e a t t h e p o i n t s o f growth. It  i s of i n t e r e s t  t o note t h a t r a d i a l growth i s more c l o s e l y  a s s o c i a t e d w i t h t h e amount o f f o l i a g e i n the 2.0-foot  interval  immed-  i a t e l y above t h e p o i n t o f growth than i t i s w i t h the t o t a l amount o f f o l i a g e above t h e p o i n t o f growth.  The o p p o s i t e o f t h i s a s s o c i a t i o n i s  t r u e o f c r o s s - s e c t i o n a l a r e a and s e c t i o n volume growth. undoubtedly  r e f l e c t s the fact  that the cumulative  This difference  f o l i a g e weight i s  p r o b a b l y as much a f u n c t i o n o f t r e e s i z e as i t i s o f p o s i t i o n w i t h i n the tree. From t h e p r e c e d i n g a n a l y s i s i t i s o b v i o u s  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 p a t t e r n one must f i r s t v a r i a t i o n s due t o t r e e s i z e .  account f o r  The f o l l o w i n g t h r e e e q u a t i o n s  appear t o be  b e s t 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 a r e a and s e c t i o n volume growth in  e n t i r e stems t o t h e q u a n t i t y o f f o l i a g e . L o g i o R.Gr.  =  2.095+0.0346 R  Logio  SVGr.  =  first  n  e q u a t i o n accounted  growth, the second  2  - 0.86  2  = 0.523  =  Logio n  2.122 + 0.183 R  The  = 0.46  C.S.A.Gr. = 0.114 + 0.598 R  Logio  2  Logio F + 0.054 D  Log  1 0  n  =  358 CFW + 0.033 D 358  CFW + 0.076 D =  358  f o r o n l y 46 p e r c e n t o f t h e v a r i a t i o n i n r a d i a l  e q u a t i o n accounted  f o r 86 p e r c e n t o f 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 a r e a growth, and the t h i r d e q u a t i o n accounted  f o r 52 p e r -  c e n t o f t h e v a r i a t i o n i n s e c t i o n volume growth. T a b l e 26 p r e s e n t s some s i m p l e 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 o f t h e r a d i a l , c r o s s - s e c t i o n a l a r e a , and s e c t i o n volume growth of  crown-formed wood and c l e a r bole-formed  wood, and s e v e r a l t r e e and  69  crown c h a r a c t e r i s t i c s .  T a b l e 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 o f r a d i a l , c r o s s - s e c t i o n a l a r e a , and s e c t i o n volume growth i n crown-formed wood and between 113 measurements o f r a d i a l , c r o s s - s e c t i o n a l a r e a , and s e c t i o n volume growth i n bole-formed wood, and s e v e r a l t r e e and s e c t i o n c h a r a c t e r i s t i c s . Correlation  Characteristic  Crown-formed wood R. Gr.  Tree height Dbh Section height S e c t i o n diameter S e c t i o n age F o l i a g e weight i n 2 f t . above Cumulative f o l i a g e w e i g h t above H e i g h t from top Percent of t o t a l height Form  p.05  The  0.620 0.744 •-0.126 0.892 0.494  (F)  0.495  0.247  0.218  (CFW) 0.290 (HT) -0.117  0.910 0.670  0.912 0.706  0. 714 0. 223  0.866 0.494  0.820 0.338  0.228 (PH) (d/D) -0.263  -0.527 0.480  •-0.570 0.541  -0. 040 0. 092  -0.236 0.271  -0.070 0.044  r  l j X 1 1  = 0.187  also  there i s l i t t l e  volume a r e a growth.  the t r e n d s suggested  a r e a and s e c t i o n  —  following  0.589 0.753 0.024 0.669 0.096 —  0.126  =  agree w i t h  those p r e s e n t e d i n  advantage i n d i f f e r e n t i a t i n g  The r e s u l t s  cross-  i n T a b l e 26 tend  e a r l i e r i n t h i s study w h e r e i n r a d i a l of p o s i t i o n w i t h i n  the t r e e whereas  volume growth i s n o t o n l y a f u n c t i o n  o f the s i z e o f t h e t r e e The  i*3  i n T a b l e 26 g e n e r a l l y  growth appears t o be a f u n c t i o n  but  2  —  0.604 0.792 •-0.150 0.835 0.289  and crown-formed wood when s t u d y i n g r a d i a l  sectional, or section  sectional  v1  0. 406 0. 621 0.027 0. 585 0. 108  C.S.A. Gr. S.-V.Gi  0.639 0.764 -0.075 0.872 0.451  T a b l e 25, s u g g e s t i n g t h a t  to support  S.V.Gr. R.Gr.  0.454 0.623 0.481 0.150 -0.310  presented  between b o l e - f o r m e d  C.S.A.Gr .  Bole--formed wood  (H) (D) (h) (d) (A)  signif. level  results  coefficients (r)  a t the p o i n t  equations  cross-  of position  where the growth i s measured.  appear t o 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 t h e r a d i a l and c r o s s - s e c t i o n a l  a r e a growth o f 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  Log 1 0 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 earlywood, 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 i n foliage distribution were not reflected in  71  i n e i t h e r earlywood  o r latewood  formation.  Conclusions  The  r e s u l t s o f the analyses i n d i c a t e that r a d i a l , c r o s s -  s e c t i o n a l a r e a , and s e c t i o n volume growth a r e h i g h l y v a r i a b l e both w i t h i n and  among 1 0 0 - y e a r - o l d  have a f f e c t e d  lodgepole pine trees.  the c u r r e n t p a t t e r n o f r a d i a l  T h i n n i n g does n o t appear t o growth ( T a b l e 2 4 ) . T h i s i s  p r o b a b l y due to t h e l a t e age a t which the stand was t h i n n e d . Kozlowski  (1960) suggested  t h a t i n o l d e r s t a n d s , wherein  Kramer and  the r e s i d u a l trees  are s m a l l crowned, i n c r e a s e s i n p h o t o s y n t h e s i s a r e accompanied by i n c r e a s e s i n r e s p i r a t i o n and the n e t g a i n i s n e g l i g i b l e . needles  I n a d d i t i o n , any shade  p r e s e n t i n the crowns o f the r e s i d u a l t r e e s were p r o b a b l y  when exposed t o f u l l s u n l i g h t f o l l o w i n g t h i n n i n g . to d e c r e a s e  from  appears  the apex towards the bottom o f the t r e e w i t h a s l i g h t  i n c r e a s e a t the base c o r r e s p o n d i n g attributed  R a d i a l growth  injured  t o the b u t t s w e l l .  t h i s phenomenon, a t l e a s t  Onaka (1950a)  i n p a r t , t o the d i s t r i b u t i o n o f  hormonal and food s u b s t a n c e s , which a r e produced  h i g h i n the crown,  d i m i n i s h i n q u a n t i t y as they move down the crown, and accumulate a t t h e base.  Because t h e t r e e s examined i n t h e p r e s e n t study cannot  suppressed,  be c o n s i d e r e d  t h e r e s u l t s o b t a i n e d a r e n o t i n complete agreement w i t h the  general patterns of r a d i a l  growth r e p o r t e d i n t h e l i t e r a t u r e  (Onaka, 1950a;  F a r r a r , 1961; L a r s o n , 1963; S h i n o z a k i et_ al_., 1964) which suggest r a d i a l growth i n c r e a s e s from the t o p to a p p r o x i m a t e l y f o l i a g e , whereafter  i t remains c o n s t a n t o r d e c r e a s e s  that  the p o i n t o f maximum 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 w e i g h t and r a d i a l growth w i t h i n the crown ( T a b l e 26) tend t o s u p p o r t H a l l ' s  (1965) and Heger's (1965) c o n c l u s i o n s  t h a t some f a c t o r s i n a d d i t i o n to n u t r i t i o n a l g r a d i e n t s determine the  72  p a t t e r n o f stem wood increment. the d i s t r i b u t i o n o f growth was a l t h o u g h t h i s d i s c r e p a n c y may when the p r e s e n t study was  However, u n l i k e H a l l ' s not c l o s e l y r e l a t e d  be due  carried  (1965) r e s u l t s ,  to d i s t a n c e from apex  to the advanced age of the t r e e s  out.  The  two  complete  stem a n a l y s e s  c a r r i e d out i n the p r e s e n t study i n d i c a t e d d i s t r i b u t i o n s o f earlywood latewood  similar  to those r e p o r t e d by Heger  (1965).  The r e s u l t s p r e s e n t e d i n F i g u r e 35, which suggest of t i s s u e produced thus crown s i z e ) ,  t h a t the amount  per pound o f f o l i a g e i s u n r e l a t e d to t r e e s i z e agree w i t h the r e s u l t s p r e s e n t e d  r e s i n o s a A i t . ) by H a l l  and  (and  f o r red p i n e ( P i n u s  (1965) but d i s a g r e e w i t h those p r e s e n t e d f o r b l a c k  spruce  ( P i c e a mariana  balsam  f i r (Abies balsamea (L.) M i l l . ) by B a s k e r v i l l e  balsam  f i r and b l a c k s p r u c e a r e r e l a t i v e l y more shade t o l e r a n t than l o d g e -  p o l e and  ( M i l l . ) BSP.)  by Weetman and H a r l a n d  (1964) and f o r  (1965a). » Because  r e d p i n e , s m a l l crowned t r e e s o f the former s p e c i e s p r o b a b l y have  a h i g h e r p r o p o r t i o n o f shade n e e d l e s which a r e p h o t o s y n t h e t i c a l l y more efficient  a t low l i g h t  insufficiently  tested,  intensities  (Kramer and K o z l o w s k i , 1960).  the r e s u l t s p r e s e n t e d i n F i g u r e 34 suggest  r e l i a b l e p r o j e c t i o n s of needle dry-weights  o f stands and  a b l e from e s t i m a t e s o f p e r i o d i c a n n u a l volume  Although that  t r e e s are o b t a i n -  increment.  A l t h o u g h 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 i m p l i c a t i o n s o f these r e s u l t s a r e unclear..  For example, a l t h o u g h c r o s s - s e c t i o n a l a r e a growth was with cumulative (stem) s i z e . n o t be used  f o l i a g e weight,  t h i s may  highly  correlated  merely be a r e f l e c t i o n o f  tree  T h e r e f o r e , the r e s u l t s p r e s e n t e d h e r e i n , by themselves, to defend or r e f u t e any  i n t r o d u c t i o n to t h i s c h a p t e r .  can-  of the form t h e o r i e s p r e s e n t e d i n the  F u r t h e r , these r e s u l t s are o f l i t t l e  i n d e f i n i n g t h i n n i n g or p r u n i n g p r e s c r i p t i o n s .  value  73  Recognizing these limitations i n the present study, i t i s possible to suggest how a study, having similar objectives, should be undertaken.  The f i r s t recommendation would be to limit the number of  measurements taken i n 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 i n planning the management of stands on the rotations likely to be used i n 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 N o r t h America,  the s o l e s o u r c e o f f i b r e f o r  p u l p i n g has been the b a r k - f r e e 'merchantable' such as t h a t r e p o r t e d by C r o s s l e y (1938) and  bole.  H i s t o r i c a l l y , research,  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 o t h e r components f o r p u l p f i b r e has been r a t h e r limited. within  However, i f the p r o j e c t e d w o r l d demand f o r w o o d - f i b r e  the next 30 y e a r s i s c o r r e c t , a s h o r t a g e o f as much as 100  c u b i c metres ( o r 44 m i l l i o n oven-dry of 25 l b . / f t . ) 3  Hatton  products  t o n s , based on an average b u l k d e n s i t y  o f wood has been f o r e c a s t by Keays (1970) and Keays and  (1971a).  In a n t i c i p a t i o n o f t h i s s h o r t a g e i n c r e a s e d a t t e n t i o n  r e c e n t l y been devoted t r e e s and  million  to c l o s e r and more complete  u t i l i z a t i o n of  has  forest  stands. To d a t e the most comprehensive  studies of complete-tree  utiliza-  t i o n have been c a r r i e d o u t by P r o f e s s o r H a r o l d Young and h i s c o l l e a g u e s a t the U n i v e r s i t y o f Maine (Chase e t a l . ,  1971;  Dyer,  Young, 1964,  1968;  Young and Chase, 1965;  and Guinn,  1965a, 1965b, 1966,  1966;  Young e t a l . ,  1967,  1964;  Young e t a l . ,  1967;  1965).  Dyer e t a l . ,  The  1968;  Young  feasibility  o f c o m p l e t e - t r e e u t i l i z a t i o n of Canadian wood s p e c i e s i s b e i n g s t u d i e d the Canadian  by  F o r e s t r y S e r v i c e a t i t s Western F o r e s t P r o d u c t s L a b o r a t o r y  (Keays, 1971a,b; Keays and H a t t o n , 1971a,b; H a t t o n and Keays, H a t t o n and Samkova, 1972).  1971;  The r e s u l t s o f these s t u d i e s suggest t h a t  those  t r e e components p r e v i o u s l y c o n s i d e r e d n o n - u t i l i z a b l e and l e f t as r e s i d u e  75  f o l l o w i n g l o g g i n g can be c o n v e r t e d  i n t o s a t i s f a c t o r y pulp.  T a b l e 27  p r e s e n t s a g e n e r a l summary o f the p u l p y i e l d and s t r e n g t h q u a l i t y o f the various previously unutilized  T a b l e 27.  t r e e components.  Comparison o f r e l a t i v e v a l u e s o f k r a f t from c o n i f e r t r e e components expressed of the b o l e .  Pulp p r o p e r t i e s  Non-merch. top (>1 i n . )  Y i e l d a t 20 permanganate no. Burst f a c t o r Tear f a c t o r Breaking length B e a t i n g time  99% ± 100% ± 85% ± 100% ± faster  Source:  Keays  Based on these e s t i m a t e s  pulp i n terms  Branches (>1 i n . )  1 5 5 5  Root + stump (>1 i n . )  70% 55% ± 5 75% ± 5 55% ± 5 faster  99% ± 1 85% ± 5 92.5% ± 2.5 85% ± 5 slower  (1971a) ( T a b l e 27) i t i s apparent  t h a t , i n terms o f p u l p  y i e l d and q u a l i t y , non-merchantable tops can and s h o u l d be u t i l i z e d . Small amounts o f branch developed Table  wood may be u t i l i z e d  i f economic means can be  t o h a r v e s t , debark, and c h i p branches.  27 suggest  pulp, u t i l i z a t i o n  the r e s u l t s i n  t h a t the root-stump component c o n v e r t s i n t o  satisfactory  i s d o u b t f u l because o f problems i n h a r v e s t i n g , c l e a n i n g ,  t r a n s p o r t i n g , debarking  and c h i p p i n g t h i s  A c c o r d i n g t o Keays (1971a), to c o m p l e t e - t r e e  Although  component.  the c r i t i c a l  q u e s t i o n s which r e l a t e  u t i l i z a t i o n are:  1.  How much o f 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 o f p u l p o r o t h e r p r o d u c t  c o u l d be o b t a i n e d  from each t r e e component? 3.  How would the v a r i o u s t r e e components be e x t r a c t e d , t r a n s p o r t e d and  processed?  76  4.  What would be the e f f e c t o f c o m p l e t e - t r e e u t i l i z a t i o n on f o r e s t growth and r e g e n e r a t i o n , on s i l v i c u l t u r a l on the t o t a l  The  purpose  first  practices,  ecology?  o f the p r e s e n t study i s t o answer, a t l e a s t  i n p a r t , the  two o f these q u e s t i o n s f o r l o d g e p o l e p i n e , a s p e c i e s o f g r e a t  importance  t o the f o r e s t economies' o f A l b e r t a and t h e I n t e r i o r o f B r i t i s h  Columbia.  Little  information pertinent  t o t h e l a t t e r two q u e s t i o n s i s  p r e s e n t l y a v a i l a b l e , and c o n s e q u e n t l y o n l y a l i m i t e d d i s c u s s i o n o f t h e s e topics i s presented.  Methods and M a t e r i a l s  Data  collection  Ten sample t r e e s from Stands Chapter  2) were used  1 and 2 ( p r e v i o u s l y d e s c r i b e d i n  to study the p u l p y i e l d  forest-grown lodgepole pine t r e e s . 1  and q u a l i t y o f 1 0 0 - y e a r - o l d ,  No attempt was made a t r a n d o m i z a t i o n  i n the s e l e c t i o n p r o c e s s , and t h e c h o i c e o f sample t r e e s was based tree's size  on t h e  (two t r e e s from each o f the 4-, 6-, 8-, 10-, and 1 2 - i n c h dbh  c l a s s e s were chosen) and apparent t r e e b o l e s i n Stand The  f r e e n e s s from i n j u r y  2 were s c a r r e d d u r i n g the e a r l i e r  t r e e s were c u t a t a 1.0-foot  thinning o f the stand).  stump, and weighed w i t h  and  foliage intact  and  f o l i a g e removed ( t o t a l stem f r e s h - w e i g h t ) .  the merchantable  ( i . e . , some o f the  branches  ( t o t a l above-ground f r e s h - w e i g h t ) and w i t h the branches The stem was s u b d i v i d e d i n t o  b o l e ( e q u a l t o o r g r e a t e r than 4.0 i n c h e s i n diameter  The c o m p l e t e - t r e e u t i l i z a t i o n p o t e n t i a l o f seven white spruce t r e e s grown i n Stand 2 were r e p o r t e d by Keays and Hatton (1971b).  (ob)),  77  non-merchantable  top ( t h a t p o r t i o n o f the stem l e s s than 4.0  g r e a t e r than o r e q u a l to 1.0 of the stem l e s s  than 1.0  inch i n diameter  (ob)) , and  inches but  t i p (that portion .  i n c h i n d i a m e t e r (ob)) , and the weights o f these  t h r e e components were measured.  The merchantable b o l e and  non-merchantable  top were each f u r t h e r s u b d i v i d e d i n t o f o u r s e c t i o n s o f e q u a l l e n g t h ,  and  o n e - i n c h t h i c k d i s c s , f o r m o i s t u r e c o n t e n t d e t e r m i n a t i o n s (as p r e v i o u s l y outlined  i n Chapter 2) were sawn from  wood samples  the end o f each s e c t i o n .  Unbarked  f o r the pulp e v a l u a t i o n s t u d i e s were o b t a i n e d from each  merchantable b o l e and non-merchantable 1.  top s e c t i o n as  follows:  For those s e c t i o n s w e i g h i n g l e s s than o r a p p r o x i m a t e l y e q u a l t o 15.0  pounds ( f r e s h - w e i g h t ) the e n t i r e s e c t i o n was  taken,  and 2.  For those s e c t i o n s w e i g h i n g more than 15.0 weight)  pounds  a sample, w e i g h i n g a p p r o x i m a t e l y 15.0  (fresh-  pounds and  sawn a t an a n g l e of 40° from a p l a n e p e r p e n d i c u l a r to the t r e e l e n g t h , was  removed from the c e n t r e of each  section.  A more d e t a i l e d d e s c r i p t i o n o f the p u l p sample c o l l e c t i o n p r o c e d u r e i s g i v e n by Keays  (1968).  The branches  and n e e d l e s were s u b d i v i d e d i n t o branches 1.0  i n d i a m e t e r and g r e a t e r , and p l u s n e e d l e s , and weighed. were c o l l e c t e d e q u a l to 1.0  less  than 1.0  A l l o f the branches  i n c h i n d i a m e t e r were r e t a i n e d because  pulp e v a l u a t i o n s t u d y .  less  than the 60.0  g r e a t e r than o r  i n a l l c a s e s the weight  pounds per t r e e d e s i r e d  2.  f o r the  A l l o f the n e e d l e - b e a r i n g twigs were c l i p p e d  l e s s than 1.0  inch i n diameter  from  (which were then d i s c a r d e d ) ,  put i n t o b u r l a p sacks and d r i e d under c o n d i t i o n s s i m i l a r i n Chapter  i n c h i n diameter  D i s c s , f o r moisture content determination,  from the b r a n c h e s .  of these branches was  the branches  i n t o branches  inch  to those noted  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 i n 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 i n 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 i n crates and shipped to the Western Forest Products Laboratory. Some of the characteristics of the ten trees used i n the pulp study are presented i n 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 nonmerchantable top),branches 1.0 inch i n 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 No. (yr.) (in.)  Height (ft.)  Merch. Non-merch. bole bole (4"-l M ) Root-St. length length < 1" (ft.) (ft.)  Component dry-weights (lb.) bark included Top Top Branch Branch Root-St. Bole > 1" > 4" 4 , , -l" •<1" > 1" < 1"  Dead Needles Branches  7.1 12.0 4.5 5.8 5.6 9.9 10.1 8.0 12.0 4.3  64.2 74.3 55.0 53.2 52.3 66.7 72.0 58.7 81.2 47.1  42.4 63.8 14.0 30.4 30.7 53.7 55.8 46.2 67.9 9.6  18.7 8.0 37.4 18.4 18.9 10.3 13.2 8.8 9.8 35.0  3.6 42.6 2.7 4.5 4.3 28.1 13.3 11.3 20.9 0.6  50.5 255.6 15.8 30.3 27.5 142.1 142.7 70.4 204.4 15.5  192.7 823.8 49.3 125.2 113.5 493.1 470.6 286.8 810.0 30.5  20.5 8.6 61.7 33.3 30.6 18.2 16.1 11.1 8.6 55.8  1.0 0.5 1.0 0.5 0.5 0.4 0.4 0.4 0.4 0.6  0.0 30.9 0.0 0.0 0.0 4.7 4.4 3.8 17.2 0.0  15.1 121.9 9.0 7.5 8.7 67.7 48.5 36.2 82.2 3.2  3.0 37.0 1.0 1.0 2.0 2.0 8.0 1.0 23.0 1.0  8.5 60.6 8.6 7.4 8.9 38.3 33.2 23.8 28.1 2.6  96. 3 79.3 Mean St.dev. 2.0 2.9  62.5 11.0  41.5 20.0  17.9 10.5  13.2 13.6  95.5 85.7  339.6 297.7  26.5 19.0  0.6 0.2  6.1 10.2  40.0 39.9  7.9 12.3  22.0 18.4  1 2 3 4 5 6 7 8 10 11  95 95 98 97 99 96 96 98 92 97  VO  80  the c h i p s were r e p r e s e n t a t i v e of the f u l l b o l e o f each t r e e , p r o r a t e d amounts o f c h i p s (based  on s e c t i o n volume w i t h i n each t r e e ) , from each  sample s e c t i o n , were t h o r o u g h l y blended t o g e t h e r . l i c a t e chip moisture  P r i o r to c o o k i n g  c o n t e n t d e t e r m i n a t i o n s , u s i n g about 60  dry c h i p s per d e t e r m i n a t i o n , were c a r r i e d out f o r each The  dup-  grams o f a i r -  tree.  l a b o r a t o r y p u l p i n g 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 r e s e a r c h d i g e s t e r w i t h a 1.0-cubic-foot c a p a c i t y . The m a t e r i a l to be pulped was  s e a l e d i n s t a i n l e s s s t e e l bombs (each o f  735-ml c a p a c i t y ) which were p l a c e d i n the r e s e a r c h d i g e s t e r . d e s c r i p t i o n o f the d i g e s t e r assembly was  (1970) .  The  experimental  presented  A  by Keays and  detailed Bagley  d e s i g n of t h i s study r e q u i r e d f o u r bombs  f o r s c r e e n i n g s , permanganate number and y i e l d d e t e r m i n a t i o n s , and f o r PFI m i l l and  strength determinations)  (two two  from each component of each  Only f i v e o f the ten sample t r e e s c o n t a i n e d branches g r e a t e r than 1.0 i n diameter.  In a d d i t i o n , i n o r d e r  f o r s c r e e n i n g s , permanganate  f o r PFI m i l l and s t e n g t h )  b o l e sample s e c t i o n s were cooked f o r one 60  grams o f a i r - d r i e d c h i p s .  inch  to e v a l u a t e the v a r i a t i o n i n p u l p  q u a l i t y w i t h i n the t r e e , two bombs (one number and y i e l d , and one  tree.  tree.  from each o f the e i g h t  Each bomb c o n t a i n e d  about  Because i t i s d e s i r a b l e to compare p u l p s  of a u n i f o r m permanganate number (Keays et a l . , 1969;  H a t t o n and  Keays,  1970,) an e x p l o r a t o r y cook, u s i n g c h i p s from the b r a n c h e s , root-stump system and  p r o r a t e d b o l e o f one  i n T a b l e 29,  was  t r e e and  the p u l p i n g c o n d i t i o n s p r e s e n t e d  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 p u l p w i t h a permanganate number o f 20  from each component.  The  approximately  permanganate numbers of the pulp produced  u s i n g these c o n d i t i o n s are a l s o presented  i n Table  29.  81  T a b l e 29.  K r a f t c o o k i n g c o n d i t i o n s used E x p l o r a t o r y cooks. Full  E f f e c t i v e a l k a l i s (%) Sulfidity (%) Time t o max.temp, (min.) Time a t max.temp, (min.) Max. temp. (°C) Liquor-to-wood ratio Unbleached pulp permanganate numbers obtained  bole  f o r the  Branches  Root-stump  16.,0,17.0,18.0 25.4 135 75 170 4.5:1  16.0,17.0,18.0 25.4 135 75 170 4.5:1  19.0,20.0,21.0 25.4 135 75 170 4.5:1  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 o f the e x p l o r a t o r y cook i t was p o s s i b l e t o e s t i m a t e the e f f e c t i v e a l k a l i r e q u i r e d to^produce permanganate number o f a p p r o x i m a t e l y conditions  ( T a b l e 30) were used  T a b l e 30.  Effective alkali:  an unbleached  20 ( F i g u r e 36).  pulp w i t h a  The f o l l o w i n g c o o k i n g  f o r a l l the subsequent cooks i n t h i s  K r a f t c o o k i n g c o n d i t i o n used o b t a i n unbleached pulps w i t h numbers o f a p p r o x i m a t e l y 20 components o f 100-year-old, lodgepole pine trees.  f u l l bole root-stump branches  (%) (%)  (%)  Sulfidity Time t o max. temp. Time a t max. temp. Max. temp. Liquor-to-wood ratio  (%) (min.) (min.)  (°C)  study.  i n study t o permanganate from the t h r e e forest-grown  15.3 14.8 16.8 25.4 135 75 170 4.5  Upon c o m p l e t i o n o f t h e c o o k i n g , b l a c k l i q u o r was c o l l e c t e d from each bomb and r e t a i n e d 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 . o f each bomb were d i s i n t e g r a t e d t h o r o u g h l y washed.  The cooked c h i p s  f o r f i v e minutes and the r e s u l t i n g pulp was  The p u l p to be used  f o r y i e l d d e t e r m i n a t i o n was oven-  82  Figure36. THE RELATIONSHIP B E T W E E N NUMBER  AND  EFFECTIVE  100-YEAR-OLD  21  r  20  -  UNBLEACHED  ALKALI  LODGEPOLE  FOR  PULP  THREE  PERMANGANATE  COMPONENTS  OF  PINE.  19  18  JO  E  17  16  IS  14  13  jl. 14  IS  16  17 Effective  19  18 Alkali  !%J  20  21  83  d r i e d a t 105°C weight.  f o r a minimum o f 16 hours and had reached a' c o n s t a n t d r y -  The p u l p to be used f o r P F I m i l l  air-dried  for a similar  time p e r i o d .  d i s i n t e g r a t e d f o r f i v e minutes  and s t r e n g t h d e t e r m i n a t i o n s was  The p u l p from each bomb was  and the amount o f r e j e c t s was  re-  determined  by  w e i g h i n g the oven-dry m a t e r i a l r e t a i n e d on a 10-cut p l a t e o f a V a l l e y laboratory f l a t  screen.  The s c r e e n e d pulp from each bomb was  f o r f i v e m i n u t e s , t h o r o u g h l y mixed  and c o n d i t i o n e d  centrifuged  to a u n i f o r m m o i s t u r e  c o n t e n t o f about 8 p e r c e n t . Unbleached  permanganate numbers were determined on the screened  pulp used f o r the y i e l d d e t e r m i n a t i o n s f o l l o w i n g T a p p i Standard ( u s i n g 40 ml  o f 0.1N  KMnOu).  Both the pH and r e s i d u a l a l k a l i l i n e d by Macdonald  Each d e t e r m i n a t i o n was  T214ts-50  done i n d u p l i c a t e .  ( f o l l o w i n g methods s i m i l a r  to t h o s e o u t -  (1969)) were determined f o r the b l a c k l i q u o r  from each  bomb. The s c r e e n e d p u l p from the two bombs f o r each component  from  each t r e e t o be used f o r the s t e n g t h d e t e r m i n a t i o n s were t h o r o u g h l y combined.  T h i s m i x t u r e was  which a sample o f 24.0 particularly  grams O.D.  was  two  equal a l i q u o t s ,  withdrawn.  Each sample was  1.2.percent u n t i l  from each o f  In some i n s t a n c e s ,  i n the c a s e o f b r a n c h e s , a s m a l l e r sample was  by low y i e l d s .  adjusted  divided into  necessitated  d i s i n t e g r a t e d a t a c o n s i s t e n c y o f about  f r e e from f i b r e b u n d l e s and c l o t s .  The sample was  to a c o n s i s t e n c y o f 10 p e r c e n t and p l a c e d i n the PFI m i l l .  then The  P F I m i l l methods used were s i m i l a r to those o u t l i n e d by Standard T e s t i n g P r o c e d u r e PB-6  o f 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 o f Canada, 1962).  Two  c o n d i t i o n i n g runs were  c a r r i e d out b e f o r e the b e a t i n g o f each component. The o b j e c t i v e o f t h i s study was  to t e s t the s t r e n g t h o f the  unbleached p u l p from the d i f f e r e n t components a t a f r e e n e s s o f 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 m i l l 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 r 2 ) were determined by the methods described previously in this thesis (Chapter 2). Unlike the results presented i n the biomass section of this thesis (Chapter 2), the results presented i n this section are limited to that  portion of the components considered to be potentially utilizable  for pulping. It i s necessary therefore to redefine the components as follows:  85  1.  Merchantable b o l e :  t h a t p o r t i o n o f the stem from a 1.0-foot  stump to a A.0-inch top d i a m e t e r ( o b ) . 2.  Non-merchantable  top:  t h a t p o r t i o n o f the stem from a  4.0-  i n c h d i a m e t e r (ob) to a 1.0-inch d i a m e t e r ( o b ) . 3.  F u l l bole: top  (i.e.,  the merchantable b o l e p l u s the  non-merchantable  t h a t p o r t i o n o f the stem from a 1.0-foot stump to  a 1.0-inch top d i a m e t e r ( o b ) . 4.  Branches:  those branches g r e a t e r than o r e q u a l to 1.0  inch i n  diameter ( o b ) . 5.  Root-stump system: from ground l e v e l roots  the stump ( i . e . , t h a t p o r t i o n of the stem to 1.0  f o o t above ground l e v e l ) p l u s the  ( i . e . , t h a t p o r t i o n o f the t r e e below ground  g r e a t e r than 1.0  level)  inch i n diameter ( o b ) .  In  the f o l l o w i n g s e c t i o n o f t h i s c h a p t e r , r e f e r e n c e s to the components a r e  to  those d e f i n e d above u n l e s s o t h e r w i s e s p e c i f i e d . I n o r d e r to e l i m i n a t e e r r o r i n the comparison o f y i e l d o f  p u l p s v a r y i n g i n permanganate number ( H a t t o n and K e a y s , 1970) pulp y i e l d a t permanganate number 20 was  calculated  an a d j u s t e d  f o r a l l y i e l d data  u s i n g the f o l l o w i n g f o r m u l a : Y20  = Y• + 3 (20.0-K ) obs. obs.  where: Y 2 u  = adjusted y i e l d  to permanganate number 20  Y , = observed J y i e l d obs. ^obs  =  °b  s e r v e c  *  permanganate number  3 = component s l o p e c o n s t a n t f o r the r e l a t i o n s h i p between p u l p y i e l d and permanganate number ( F i g u r e 37) and e q u a l s 0.38,  0.36, and 0.43  f o r the  f u l l b o l e , r o o t - s t u m p , and b r a n c h components, respectively.  The 0.38  s l o p e observed f o r the  full  b o l e agrees w i t h t h a t r e p o r t e d f o r l o d g e p o l e p i n e  Permanganate Number  87  by H a t t o n and Keays (1970) .  R e s u l t s and D i s c u s s i o n  The q u a n t i t y o f each component p o t e n t i a l l y a v a i l a b l e from the completet r e e u t i l i z a t i o n o f 100-year-old l o d g e p o l e p i n e  trees  The e q u a t i o n s p r e s e n t e d i n T a b l e 6 and F i g u r e s 6, 10, and 11 are  o f use i n o b t a i n i n g gross e s t i m a t e s o f the wood r e s o u r c e p o t e n t i a l l y  a v a i l a b l e through c o m p l e t e - t r e e u t i l i z a t i o n . o b t a i n e d a r e o v e r e s t i m a t e s because ponents  However, the e s t i m a t e s thus  they i n c l u d e such n o n - u t i l i z a b l e com-  as bark and p o r t i o n s o f t h e components l e s s than 1.0  diameter.  inch i n  U n f o r t u n a t e l y , s u f f i c i e n t d a t a n e c e s s a r y t o develop  t a r y b a r k - f r e e , oven-dry weight  complemen-  e q u a t i o n s f o r the components, as r e d e -  f i n e d on page 85 o f t h i s t h e s i s , were n o t a v a i l a b l e . B a r k - f r e e oven-dry  wood weights f o r each component a r e shown  i n T a b l e 31 e x p r e s s e d as percentages o f the f u l l b o l e .  T a b l e 31.  T r e e Dbh no. ( i n . )  i  Merch. bole Wt.  175.5 762.8 45.3 115.2 103.1 455.6 436.2 261.6 754.1 27.9 313.7 Mean Stan . dev. 276.9 1 2 3 4 5 6 7 8 10 11  7.1 12.0 4.5 5.8 5.6 9.9 10.1 8.0 12.0 4.3  B a r k - f r e e , oven-dry weights o f the 100-year-old l o d g e p o l e p i n e t r e e components as a p e r c e n t a g e of f u l l b o l e s .  %  90.8 99.1 45.1 79.4 79.6 96.6 96.9 96.5 99.0 35.7 81.9 23.2  Tree components (weight i n l b . ) Branches Non-merch. Full bole bole Wt. Wt. Wt. % % %  9.2 17.7 7.1 0.9 55.2 54.9 29.8 20.6 26.4 20.4 3.4 15.8 13.8 3.1 3.5 9.4 1.0 7.4 50.3 64.3 23.3 18.1 17.3 23.2  193.2 769.9 100.5 145.0 129.5 471.4 450.0 271.0 761.5 78.2 337.0 263.8  100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0 100.0  -  0.0 24.5 0.0 0.0 0.0 3.7 3.6 2.9 12.8 0.0 4.8 8.0  0.0 3.2 0.0 0.0 0.0 0.8 0.8 1.1 1.7 0.0 0.8 1.0  Root-•Stump Wt.  43.9 231.1 14.6 27.3 24.7 129.6 127.3 63.4 187.4 13.4 86.3 78.0  % 22.7 30.0 14.5 18.8 19.1 27.5 28.3 23.4 24.6 17.7 22.6 5.1  88  F i g u r e 38  illustrates  f r e e , oven-dry weight expressed and  t r e e diameter.  the r e l a t i o n s h i p between component bark-  as a p e r c e n t a g e  o f the f u l l b o l e weight  In s e l e c t i n g an e q u a t i o n s u i t a b l e f o r d e s c r i b i n g  the  r e l a t i o n s h i p s f o r the merchantable b o l e and the non-merchantable t o p , two c o n d i t i o n s were s o u g h t , namely: a)  The  e q u a t i o n f o r the non-merchantable top s h o u l d e q u a l  p e r c e n t at a dbh l i m i t ) and  (ob) of 4.0  asympototic  100  i n c h e s (the m e r c h a n t a b i l i t y  to 0.0  p e r c e n t at l a r g e  diameters,  and b)  The  e q u a t i o n f o r the merchantable b o l e s h o u l d e q u a l  p e r c e n t at a dbh 100.0 The  (ob) o f 4.0  p e r c e n t at l a r g e  e q u a t i o n s p r e s e n t e d i n F i g u r e 38  components g r e a t e r than 1.0  s a t i s f y the above c o n d i t i o n s .  c o n d i t i o n i n g was  not attempted  were the b e s t e q u a t i o n s  ( i . e . , the  percentages  too narrow t o i n c l u d e  f o r the root-stump and b r a n c h  and  components,  the e q u a t i o n s p r e s e n t e d i n F i g u r e  ( i n terms o f s t a n d a r d e r r o r s o f e s t i m a t e  c o e f f i c i e n t s of determination)  and  38  and  obtained.  As can be seen from the r e s u l t s p r e s e n t e d i n T a b l e 31 38  and  f o r s m a l l t r e e s t o a p r o b a b l e maxima  f o r l a r g e t r e e s ) the range o f t r e e s i z e s samples was Consequently,  Although  a p p l y to the root-stump  i n c h i n diameter  f o r these components i n c r e a s e from 0.0  these c o n d i t i o n s .  a s y m p t o t i c to  diameters.  s i m i l a r t h e o r e t i c a l c o n d i t i o n s undoubtedly branch  i n c h e s and  0.0  and F i g u r e  the q u a n t i t y o f wood f i b r e c o n t a i n e d i n the non-merchantable t o p , root-stump system i s c o n s i d e r a b l e .  branches,  Because p r e s e n t h a r v e s t i n g 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 o f the non-merchantable t o p , t h i s component i s o f immediate i n t e r e s t . c o n f i r m those r e p o r t e d by Smithers  The  r e s u l t s presented i n Table  32  (1961) and Lee (1967), and demonstrate  t h a t , even i n o l d e r aged stands over a wide range o f s i t e q u a l i t i e s ,  the  Figure 38. THE RELATIONSHIP BETWEEN COMPONENT  WEIGHT  DRX BARK-FREE  TREE SIZE AND  O V E N - D R Y , BARK-FREE  EXPRESSED AS A PERCENTAGE  FULL BOLE WEIGHT  OF THE  FOR 100-YEAR-OLD  OVEN-  LODGEPOLE  PINE TREES.  loo  | ( )  NMT  [%]  OBH  « 4.255-  [in.]  3.884  log  08H  - 0.01668 tog  [DBH-  3.99°3  90  numbers of lodgepole pine trees per acre present i n the 4- to 8-inch diameter classes are substantial.  It i s apparent from the results presented  in Figure 38 that the percentage of the f u l l bole contained in the nonmerchantable 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 i n diameter classes less than 9.0 inches dbhob from eighty-eight stands of 100-year-old lodgepole pine. Dbhob Class (in.) 1  2  3  4  5  6  7  8  Total  Total  <9.0 i n .  1.0 in,  34 100 172 171 148 Ave. : 20 66 107 819 884 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 i s small (Figure 38). According to Smithers (1961), the occurrence of large branches on lodgepole pine trees i s mainly limited to trees grown i n the open or in low-density stands (100 to 600 mature trees per acre).  Relatively  l i t t l e i s known about the rooting habit of lodgepole pine trees.  Horton  (1958) reported changes i n root development at different ages and on different s o i l types. Because there i s 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 i n 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-yearold 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 100year-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  T a b l e 33.  Component pulp y i e l d d a t a from k r a f t p u l p i n g f o r t e n , 100-year-old l o d g e p o l e p i n e t r e e s . Unadjusted unscreened y i e l d (%)  Component:  Permanganate number  Adjusted unscreened y i e l d (%)  Screenings %  F u l l bole:  Mean  Range 95% C . I . (%-width) Reference  standard  1  47.11  20.31  46.99  45.54-48.86  18.8-23.3  45.33-48.78  0.45  0.59  0.52  44.3  18.5  0.84  0.11-2.20 0.31  44.8  Root-stump:  Mean Range 95% C.I. (%-width)  46.00 41.26-47.46  19.82 18.60-21.10  46.06 41.76-47.50  0.76  0.33  0.72  38.85 37.68-39.93  20.49 19.60-21.80  38.64 37.50-39.84  0.58  0.54  0.65  0.64 0.16-1.78 0.21  Branches:  2  Mean Range 95% C . I . (J^-width)  0.05 0.00-0.20 0.05  (1970) and Keays (1972).  1  H a t t o n and Keays  2  Branches a v a i l a b l e from 5 t r e e s o n l y .  The average a d j u s t e d pulp y i e l d o f 46.99 p e r c e n t a t 20 permanganate number for of  t h e f u l l b o l e component  i s somewhat h i g h e r than the a d j u s t e d pulp  yield  44.8 o b t a i n e d f o r l o d g e p o l e p i n e t r e e s o f a s i m i l a r age by H a t t o n and  Keays (1970). Values o f b u r s t f a c t o r , b r e a k i n g l e n g t h , t e a r f a c t o r , and b u l k at old  300 ml CSF o f p u l p handsheets prepared  from the components o f 100-year-  l o d g e p o l e p i n e t r e e s a r e summarized i n T a b l e 34.  93  T a b l e 34.  Component p u l p q u a l i t y a t 300 ml CSF from k r a f t p u l p i n g f o r t e n , 100-year-old l o d g e p o l e pine trees. Unbleached p u l p  Component:  quality  Burst factor  Breaking l e n g t h (m)  Tear factor  Bulk (cc/gm)  97.2  12,648 12,150-13,480 358  115 104-125 5.3  1.41 1.38-1.44 0.02  14,400±350  105±4  1.41+0.02  9,952 9,080-10,990 485  120 103-143 9.0  1.38 1.31-1.45 0.03  7,002 6,726-7,450 361  103 100-107 3.4  1.28 1.25-1.29 0.02  F u l l bole: Mean  93.0-104.0  Range 95% C.I.  2.9  (%-width)  Reference  standard  1  106±1  Root-stump: 74.3  Mean  67.1-83.2  Range 95% C.I.  (Js-width)  3.4  Branches: 53.2  Mean  51.6-55.0  Range 95% C . I .  1  (%-width)  1.6  Keays (1972). Branches a v a i l a b l e from 5 t r e e s o n l y .  Because o f d i f f e r e n c e s i n b e a t i n g t e c h n i q u e s was  used  ( i . e . , a PFI  mill  i n t h i s study and a V a l l e y b e a t e r was used by Keays (1972)) the  pulp q u a l i t y r e s u l t s presented  i n T a b l e 34 are not d i r e c t l y  comparable.  However, t h e s i m i l a r i t i e s between t h e r e s u l t s o b t a i n e d and the r e p o r t e d by Keays (1972) a r e s u f f i c i e n t l y c l o s e t o suggest  results  t h a t the  results  o b t a i n e d a r e p r o b a b l y r e p r e s e n t a t i v e o f the s p e c i e s . The b o l e pulp to a lower  lower y i e l d  and s t r e n g t h o f branchwood pulp compared to f u l l  ( T a b l e s 33 and 34), as Keays (1971d) p o i n t e d o u t , i s p r o b a b l y due c e l l u l o s e c o n t e n t , and h i g h e r l i g n i n , mannan, pentosan  c o n t e n t o f the branchwood.  and a s h  These d i f f e r e n c e s a r e a m a n i f e s t a t i o n o f t h e  94  h i g h e r percentage similarity  o f compression  wood i n the b r a n c h e s .  The g r e a t e r  i n p u l p y i e l d and q u a l i t y between t h e root-stump and f u l l  bole  components than between t h e branches and t h e f u l l b o l e i s p r o b a b l y a r e l e c t i o n o f the g r e a t e r s i m i l a r i t i e s  i n specific  gravity, f i b r e length  and w i d t h , and c e l l w a l l t h i c k n e s s between root-stumpwood and bolewood (Keays,  1971f). T a b l e 35 p r e s e n t s t h e s i m p l e 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 t r e e d i a m e t e r  (D) and a d j u s t e d pulp y i e l d  s e v e r a l components o f 1 0 0 - y e a r - o l d  and p u l p q u a l i t y f o r  l o d g e p o l e p i n e cooked by t h e k r a f t  process.  T a b l e 35.  Component  F u l l bole:  Root-stump:  Branches:3  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 t r e e s i z e (D) and component unscreened p u l p y i e l d ( a t permanganate number 20) and component unbleached p u l p q u a l i t y ( a t 300 ml CSF) from k r a f t p u l p i n g t e n , 100-year-old lodgepole pine t r e e s .  Independent variable  D D D D D D D D D  2  3  2  3  2  3  Adjusted yield1  Dependent v a r i a b l e Breaking Burst Tear 2 l e n g t h factor factor2  Bulk*  0.822 0.815 0.795  -0.279 -0.332 -0.369  -0.547 -0.559 -0.563  0.245 0.261 0.262  0.384 0.396 0.391  -0.006 0.057 0.108  0.199 0.201 0.203  0.092 0.095 0.098  -0.005 0.015 0.040  0.050 0.110 0.164  -0.368 -0.338 -0.310  0.089 0.091 0.093  0.500 0.507 0.511  -0.823 -0.804 -0.783  -0.709 -0.713 -0.714  Two o b s e r v a t i o n s p e r t r e e (p.05 s i g n i f . l e v e l r i > 1 8 = 0.444) One o b s e r v a t i o n per t r e e (p.05 s i g n i f . l e v e l r i ^ s 3 0.623) Branches a v a i l a b l e from 5 t r e e s o n l y (p.05 s i g n i f . l e v e l r j . } 3 = 0.878)  95  As can be seen from the r e s u l t s p r e s e n t e d r e l a t i o n s h i p between the a d j u s t e d p u l p y i e l d t r e e s i z e was s i g n i f i c a n t . of  i n T a b l e 35 o n l y the  o f t h e f u l l b o l e component and  The r e l a t i o n s h i p between the a d j u s t e d pulp  t h e f u l l b o l e component and t r e e s i z e appears to be o f a l i n e a r  ( T a b l e 35) and i s p r e s e n t e d statistically significant tree size expect  i n F i g u r e 39.  Although  form  t h e r e i s a p o s i t i v e and  r e l a t i o n s h i p between f u l l b o l e pulp y i e l d and  ( F i g u r e 3 9 ) , t h e s l o p e o f the r e l a t i o n s h i p i s s m a l l and one can  l i t t l e r e l a t i v e change i n f u l l b o l e p u l p y i e l d  range o f d i a m e t e r c l a s s e s . presented  yield  over  T h i s c o n c l u s i o n i s supported  a f a i r l y wide  by t h e r e s u l t s  i n T a b l e 36 which i n d i c a t e t h a t t h e r e 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 d i a m e t e r c l a s s e s b u t t h a t t h e r e were s i g n i f i c a n t  differ-  ences between t r e e s w i t h i n d i a m e t e r c l a s s e s .  T a b l e 36.  A n a l y s i s of v a r i a n c e of d u p l i c a t e determinations o f f u l l b o l e pulp y i e l d f o r two t r e e s i n each o f f i v e d i a m e t e r c l a s s e s o f 100-year-old l o d g e p o l e p i n e t r e e s .  Source  Among diam. c l a s s e s Between t r e e s w i t h i n diam. c l a s s e s Error Total  p.05  significance level  df  Mean square  4  3.73  2.79  5 10 19  1.34 0.15 1.22  8.67  F s ^ i r ^ 3.33,  ^ j S  1 3  F  5.19  These r e s u l t s g e n e r a l l y c o n f i r m t h e r e s u l t s r e p o r t e d by C h i d e s t e r e t a l . , (1939) which i n d i c a t e d t h a t the y i e l d varied did  l i t t l e when c o n s i d e r e d  on a w e i g h t b a s i s .  of jack pine  pulp  C h i d e s t e r e t a l . , (1939)  r e p o r t , however, t h a t when c o n s i d e r e d on a volume b a s i s , because o f t h e  h i g h e r bulk d e n s i t y o f slower  growing t r e e s , k r a f t pulp y i e l d s were  Figure 3 9 . T H E R E L A T I O N S H I P TREE  SIZE.  BETWEEN  TOTAL  UNSCREENED  PULP YIELD  AND  97  appreciably higher for the smaller sizes than for the larger sizes. Although tree size or growth rate did appear to affect the stength properties of jack pine pulp (Chidester e_t al_. , 1939) , i t s influence was confounded by variation resulting from position within the tree and no clearcut 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 i n 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  T a b l e 37.  General d i f f e r e n c e s i n the wood and p u l p c h a r a c t e r i s t i c s o f non-merchantable tops compared w i t h merchantable b o l e s f o r coniferous species. 1  Characteristic  Non-merch. top r e l a t i v e to merch. b o l e  Known g e n e r a l wood o r f i b r e  characteristics:  Specific gravity Fibre length C e l l wall thickness Wind throw Percentage knots P e r c e n t a g e r e a c t i o n wood Percentage l i g n i n Percentage a l p h a - c e l l u l o s e Pulp c h a r a c t e r i s t i c s characteristics):  ( 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  Weight y i e l d Tear f a c t o r Burst factor Tensile strength B e a t i n g time P e r c e n t k n o t t e r and s c r e e n r e j e c t s  1  Source:  Lower Shorter Thinner Higher Higher Higher Higher Lower  Lower Lower Higher Higher Faster Higher  Keays (1971b)  Study o f the w i t h i n - t r e e v a r i a t i o n s i n p u l p y i e l d was l i m i t e d  to u n r e p l i c a t e d t e s t s o f p u l p from e i g h t d i f f e r e n t  w i t h i n one sample t r e e as o u t l i n e d on page 77 o f t h i s t h e s i s . and  and q u a l i t y locations T a b l e 38  F i g u r e s 40 to 44 i l l u s t r a t e both t h e v a r i a t i o n w i t h i n t h e t r e e and  the d i f f e r e n c e s between the merchantable  b o l e 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 Q u a r t e r B ~ S«cond Quortar C - Third  Qvarhrr  D - Top Quart**  N o n - 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.  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  Quorter  A - Bottom  8 - Second  Quarter  B - Second  C - Third  Quarter  C - Third  0 - Top Q u a r t e r  N o n *~March Top  Figure 44. BULK  FACTOR (AT 300m! CSF) FOR UNBLEACHED  KRAFT PULP FROM SEVERAL LOCATIONS WITHIN  KRAFT PULP FROM SEVERAL LOCATIONS WITHIN  A 100-YEAR-OLD LODGEPOLE PINE TREE.  A 1O0-YEAR-OLD  A - Bottom  Quart**  B ~ Second C - Third  LODGEPOLE PINE TREE.  A - Bottom  Quarter  B ~ Second  Quorter  C  D - Top Q u a r t e r  S  Quart<  Quarter  D - Top Q u a r t e r  N o n " M e r c h . Top  Figure 43. BREAKING LENGTH (AT 300 ml CSF) FOR UNBLEACHED  Quarter  - Third  Quorter Quarter Quarter  0 ~ Top Q u a r t e t  13000  i A i N o n - M e r c h . Top  B  ,  C  M e r c h . Bole  D , , A i  B  l l ,  C  N o n - M e r c h . Top  D | 1  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  Non-merchantable top Merchantable bole Full bole  Adjusted unscreened yield1 (%)  Pulp strength  at 300 ml CSF  Burst factor  Breaking Tear length (m) factor  46.01 47.20 47.00  103.0 104.2 104.0  13,751 13,473 13,524  98 112 110  Bulk (cc/gm) 1.36 1.41 1.40  Prorated averages of observations at four different locations in the non-merchantable top and four locations i n the merchantable bole.  With the exception of burst factor, the results presented i n Table 38 were as expected. As can be seen from the results i n 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 i n 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.  102  i  Conclusions  Some of the methods used i n 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 i n 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 m i l l 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 f a c i l i t a t i n g the use of bombs for cooking.  This i n  turn greatly increases the f l e x i b i l i t y 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, indicated 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 m i l l 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-utilizable i s substantial (Figure 38).  Of those presently unutilized com-  ponents, the non-merchantable top i s of immediate interest because: a.  The harvesting and hauling of this component can be easily accommodated by present ( f u l l 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 i s substantial (Table 32) and as i s demonstrated by the results presented i n Figure 38 the percentage of the f u l l bole contained i n the non-merchantable top within this diameter range varies from 100.0 to 5.5 percent.  It i s apparent therefore, that  the yield of wood fibre per acre could be greatly increased by harvesting to a 1.0-inch top diameter.  As can be seen from the results  presented for the f u l l bole in Table 38, pulp from this component i n combination with pulp from the merchantable bole i s 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 i n combination with the merchantable bole ( i . e . , the f u 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 i n 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  f e r t i l i z a t i o n , 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 i n specific gravity, the results obtained suggest that a substantial increase i n 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 i n 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 f u l l bole.  Commercial u t 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 i n combination with f u l l 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 proportion of the total tree (approximately 20 percent). Despite the relatively high yield and quality of pulp obtainable from this component (Tables 33 and 34), commercial utilization of this component, in the foreseeable future, i s doubtful for the following reasons: a.  Extraction of the root-stump system would be d i f f i c u l t and time consuming, and, because of the bulky and irregular nature of this component, conventional hauling methods would be unsatisfactory.  b.  Major problems in removing s o i l and rock particles from this component (which should be done i n 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 s o i l erosion, s o i l 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 i n lodgepole pine trees and stands i s not presently u t i l i z e d .  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.  Utiliz-  ation of the root-stump and branch components i s 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 wildfire. reported  pine o f t e n regenerates  over-abundantly  following  Stand d e n s i t i e s o f as h i g h as 500,000 stems per a c r e have been i n young s t a n d s by Smithers  (1959) and  even as many as 100,000  l i v i n g stems per a c r e have been r e p o r t e d i n a 7 0 - y e a r - o l d s t a n d by Mason (1915). Horton old,  Under t h e s e c o n d i t i o n s c o m p e t i t i o n and  s t a g n a t i o n are s e v e r e .  (1956) r e p o r t e d a dominant h e i g h t o f o n l y 4 f e e t i n one  dense l o d g e p o l e p i n e s t a n d .  50-year-  Indeed, i t i s u n l i k e l y t h a t  stands  c o n t a i n i n g more than 2,000 stems per a c r e a t 90 years o f age w i l l yield  a r e a s o n a b l e merchantable volume ( S m i t h e r s , 1961). Remedial measures, such as t h i n n i n g o r weeding, may  s u c c e s s f u l l y undertaken  of usable m a t e r i a l .  little  o f t e n the f o r e s t manager i s f a c e d w i t h the  promise and  In o l d e r stands  p r o p o s i t i o n o f c l e a r i n g o f f the e x i s t i n g stand r e t u r n ) and production.  such measures h o l d  (with l i t t l e  expensive financial  then p l a n t i n g i f he wants to b r i n g the a r e a back i n t o  immediate  I f the m a t e r i a l s removed i n the c l e a r i n g o p e r a t i o n c o u l d be i n t o p u l p , i t would be p o s s i b l e to d e f r a y , a t  p a r t , some of the c l e a r i n g and i s t o d e s c r i b e the y i e l d  and  reforestation costs.  The  least  purpose o f  q u a l i t y of p u l p which c o u l d be  from an o v e r l y dense, h i g h l y suppressed trees.  to  or no  economically converted  study  be  i n young s t a n d s t o overcome s t a g n a t i o n and  i n c r e a s e the y i e l d  in  ever  stand o f 1 0 0 - y e a r - o l d  this  produced  lodgepole pine  108  Methods and M a t e r i a l s  Data  collection Ten sample t r e e s from Stand 3 ( d e s c r i b e d p r e v i o u s l y i n Chapter  2)  were used  to study the p u l p y i e l d  year-old lodgepole pine t r e e s .  100-  and q u a l i t y of h i g h l y suppressed  No attempt was  made to randomize the  s e l e c t i o n o f the sample t r e e s and o n l y t r e e s a p p a r e n t l y f r e e from d i s e a s e ( e . g . , dwarf m i s t l e t o e and of diameter  a t r o p e l l i s canker)*were  chosen  to cover a  range  classes. The t r e e s were cut a t a 1.0-foot stump, and weighed w i t h  and  foliage intact  ( t o t a l above-ground f r e s h - w e i g h t ) and w i t h the  and  f o l i a g e removed ( t o t a l stem f r e s h - w e i g h t ) .  branches  branches  A 1.0-inch t h i c k d i s c , f o r  m o i s t u r e c o n t e n t d e t e r m i n a t i o n s (as p r e v i o u s l y o u t l i n e d i n Chapter  2)  was  removed a t stump h e i g h t and a t 6.0-foot i n t e r v a l s above stump h e i g h t . t r e e was  d i v i d e d i n t o t h r e e components: r o o t p l u s stump, b o l e , and  plus needles.  The branches  and n e e d l e s were d r i e d  to t h o s e d e s c r i b e d p r e v i o u s l y i n Chapter  2),  oven-dry  weights  o f the b o l e and root-stump  p r e v i o u s l y d e s c r i b e d i n Chapter The  f u l l b o l e and  branches  (under s i m i l a r c o n d i t i o n s  and because none of the  were g r e a t e r than o r e q u a l to 1.0-inch i n diameter  Each  branches  they were d i s c a r d e d .  components were determined  The as  2.  the root-stump  components o f each of the t e n  t r e e s were then t r a n s p o r t e d to the Western F o r e s t P r o d u c t s Subsequently,  one o f the sample t r e e s was  component was  misplaced i n t r a n s i t .  r e j e c t e d because i t s root-stump  T a b l e 39  c h a r a c t e r i s t i c s o f the n i n e suppressed  Laboratory.  p r e s e n t s some of the  t r e e s used  f o r the pulp  study.  The e f f e c t s o f a t r o p e l l i s canker and dwarf m i s t l e t o e on p u l p y i e l d and q u a l i t y have been r e p o r t e d by Hunt and Keays (1970), and Hunt (1971), respectively.  109  Table 39.  Characteristics of the nine suppressed, 100-year-old lodgepole pine trees used i n this study.  Tree characteristic  Diameter Height Crown length Crown width Oven-dry stem weight Oven-dry root-stump weight  Mean  (in.) (ft.) (ft.) (ft.) (lb.) (lb.)  Standard dev.  2.0 22.0 12.5 3.2 20.3 8.1  0.9 5.5 4.8 1.0 16.3 6.3  Min. value  Max. value  1.2 13.2 8.0 1.5 2.1 0.6  4.5 33.7 23.7 4.0 55.5 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 i n 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  d e t e r m i n a t i o n s were made. The  c h i p s were pulped by t h e k r a f t p r o c e s s u s i n g t h e p u l p i n g  c o n d i t i o n s presented  T a b l e 40.  i n T a b l e 40.  K r a f t cooking c o n d i t i o n s f o r the bole chip mixture o f n i n e suppressed, 1 0 0 - y e a r - o l d l o d g e p o l e p i n e t r e e s .  Sulfidity Effective alkali Time t o max. temp. Time a t max. temp. Max. temp. Liquor-to-wood ratio  25.0 17.0 135.0 80.0 170.0 4:1  (%) (%) (min.) (min.) (°C)  The preceding pulping conditions (Table 40) were selected to obtain an unbleached pulp permanganate number of close to 20. The  l a b o r a t o r y p u l p i n g 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 r e s e a r c h d i g e s t e r w i t h a 1.0-cubic Upon c o m p l e t i o n  o f t h e c o o k i n g t h e p u l p was t h o r o u g h l y washed, c e n t r i f u g e d  f o r 5 minutes,  and mixed.  used  determinations p r i o r to screening.  for yield  was determined  foot capacity.  by weighing  F i v e s e p a r a t e oven-dry a l i q u o t s o f p u l p were The amount o f r e j e c t s  t h e oven-dry m a t e r i a l r e t a i n e d on a 10-cut  p l a t e of a V a l l e y laboratory f l a t  screen.  Unbleached permanganate and Kappa numbers were determined on the s c r e e n e d p u l p f o l l o w i n g T a p p i Standards respectively.  T214 ts-50 and T236/m-60,  Three d e t e r m i n a t i o n s o f each were made on t h e p u l p .  One b e a t e r r u n was done f o r the pulp u s i n g a V a l l e y l a b o r a t o r y b e a t e r f o l l o w i n g t h e method o u t l i n e d by T a p p i Standard T200 t s - 6 6 . f r e e n e s s was t e s t e d ( T a p p i Standard  Pulp  T227 Os-58) and pulp samples f o r  handsheets were c o l l e c t e d a t time i n t e r v a l s o f 0,5,25,50,60 and 72 minutes.  S i x handsheets f o r each o f the p r e v i o u s l y mentioned b e a t i n g  were prepared  a c c o r d i n g to T a p p i Standard  T205 m-58.  times  The f i v e b e s t hand-  Ill  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  Because the amount of chips available from the root-stump 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 i n 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 i n stainless steel bombs (each of 735-ml capacity) and placed i n 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 i n Table 41.  Table 41. Kraft cooking conditions for the root-stump chip mixture of nine suppressed, 100-year-old lodgepole pine trees. Sulfidity Effective a l k a l i Time to max. temp. Time at max. temp. Max. temp. Liquor-to-wood ratio  (%) (%)  (min.) (min.) (°C)  25.4 14.8 135 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  f o r s c r e e n i n g s , permanganate number, and y i e l d  were o v e n - d r i e d 105°C.  The  to a c o n s t a n t dry-weight  remaining  two  each sample was was of  determined  by w e i g h i n g  f o r PFI m i l l and s t r e n g t h  f o r a s i m i l a r time p e r i o d .  then r e d i s i n t e g r a t e d  a Valley laboratory f l a t  centrifuged  f o r a minimum of 16 hours a t  samples, to be used  d e t e r m i n a t i o n s , were a i r - d r i e d  determinations,  f o r 5 minutes and  The  pulp  from  the amount of r e j e c t s  the oven-dry m a t e r i a l r e t a i n e d on a 10-cut screen.  f o r f i v e minutes,  The  s c r e e n e d pulp from each bomb  t h o r o u g h l y mixed, and  plate was  c o n d i t i o n e d to a u n i f o r m  m o i s t u r e c o n t e n t of about 8 p e r c e n t . Unbleached permanganate numbers were determined p u l p used  f o r the y i e l d  ( u s i n g 40  ml o f 0.1  used  determinations  N KMn0i») .  The  f o l l o w i n g T a p p i Standard  screened  pulp from the two  f o r s t r e n g t h d e t e r m i n a t i o n s were t h o r o u g h l y mixed.  divided  i n t o two  (o.d.) was  1.2  Each sample was  the PFI m i l l . be s u f f i c i e n t l y the second  samples to  For the reasons  Each sample  was  o b t a i n e d a t 22,000 b e a t i n g r e v o l u t i o n s o f s t a t e d p r e v i o u s l y , t h i s was  not c a r r i e d  P u l p f r e e n e s s was  grams  p l a c e d i n the P F I m i l l .  c l o s e to the t a r g e t f r e e n e s s o f 300  PFI m i l l run was  was  d i s i n t e g r a t e d at a consistency of  then a d j u s t e d to a c o n s i s t e n c y o f 10 p e r c e n t and CSF was  ts-50  T h i s mixture  p e r c e n t u n t i l f r e e from f i b r e b u n d l e s .  A p u l p f r e e n e s s o f 306  screened  T214  equal a l i q u o t s from each o f which a sample of 24.0  withdrawn.  approximately  on the  CSF and  c o n s i d e r e d to consequently  out.  t e s t e d f o l l o w i n g T a p p i Standard  and  s i x handsheets were prepared  The  f i v e b e s t handsheets were t e s t e d f o r b r e a k i n g l e n g t h , and  T227 os-58  a c c o r d i n g t o T a p p i Standard T205  t e a r i n g s t r e n g t h f o l l o w i n g T a p p i Standard  T220  m-60.  burst  m-58. and  be  113  Analysis  Because, for the reasons outlined i n 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 J yield obs 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 i n the present study. Table 42 presents unadjusted and adjusted (to permanganate number 20) unscreened pulp yield data obtained for the suppressed trees used i n this study in comparison with similar data obtained for normally grown lodgepole pine and for the reference standard.  114  T a b l e 42.  A comparison between unscreened pulp y i e l d d a t a o f suppressed p i n e t r e e s , more n o r m a l l y grown p i n e t r e e s and the r e f e r e n c e s t a n d a r d .  Unadjusted y i e l d (%)  Permanganate number  Adjusted y i e l d (%)  Kappa no.  Screenings (%)  Bole: Supp.pine: mean range Normal pine: * 1  mean range  R e f . s t a n d : mean range  44.17 43.88-44.36  17.9  44.96 44.67-45.15  47.11 45.54-48.86  20.3 18.8-23.3  46.99 45.33-48.78  43.70  17.9  44.8  1  43.65-43.76  1  1  25.9  0.60  0.84 0.11-2.20 0.40  2  3  17.85-17.90  Root-stump: Supp.pine: mean range Normal pine: * 1  1  2  3  "*  mean range  44.86 44.40-45.31  22.0 21.4-22.6  44.14 43.90-44.37  0.63 0.20-1.06  46.00 41.26-47.46  19.8 18.6-21.1  46.06 41.76-47.50  0.64 0.16-1.78  Ref. s t a n d , from T a b l e I I I , Cook 75, H a t t o n and Keays (1970). Ref. s t a n d , from T a b l e V I , H a t t o n and Keays (1970). Ref. s t a n d , from Cook 75, u n p u b l i s h e d d a t a ( H a t t o n , 1972). T a b l e 33, Chapter 5 o f t h i s t h e s i s .  T a b l e 43 p r e s e n t s the b e a t i n g t i m e s , b u r s t f a c t o r s ,  tear  and b r e a k i n g l e n g t h s a t 500 and 300 ml CSF o f the p u l p handsheets from t h e suppressed  factors,  prepared  t r e e s i n comparison w i t h d a t a o b t a i n e d f o r more n o r m a l l y  grown p i n e and f o r t h e r e f e r e n c e s t a n d a r d .  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.  Burst factor  500 ml CSF Tear Break, factor length (m)  Burst factor  300 ml CSF Tear Break, factor length (m)  Bole: Supp.pine 94 Normal pine 1 — Ref. Stand.2 100±3  103  12,750  114 ±9  13,800±400  107 97 106±1  93 114 105±4  72.4 74.6  131.0 116.4  14,150 12,598 14,400±350  Root-stump: Supp.pine Normal pine 1 1 2  9,761 9,887  Table 34, Chapter 5 of this thesis. Keays (1972).  As demonstrated by the results presented in Table 42, the unscreened yield of pulp produced from the suppressed trees i s essentially the same as that obtained for more normally grown lodgepole pine.  Similarly,  no substantial difference was noted i n the percent screenings. The results also indicate that the quality of unbleached pulp produced from the suppressed trees is comparable to that produced from normally grown trees (Table 43).  For the f u l l 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 i n this study. However, as i s demonstrated by the results presented in Table 32, even more normally  116  grown stands  o f 100-year-old  small trees similar of pulp  lodgepole  to those  produced from these  p i n e may  c o n t a i n a l a r g e number o f  examined i n t h i s study. s u p p r e s s e d t r e e s was  t h a t produced from more n o r m a l l y  grown t r e e s ( T a b l e s 42  i s dependent upon the economics of h a r v e s t i n g and and  of f i b r e stands full  quality.  The  ( T a b l e 7 ) , and  may  be  sufficient  y i e l d and q u a l i t y  essentially  r e s u l t s s u g g e s t t h e r e f o r e t h a t commercial u t i l i z a t i o n  yield  The  and  the same as 43).  These  of t h i s wood  p r o c e s s i n g and  not  r e s u l t s presented  h e r e i n suggest t h a t the  q u a l i t y and  of p u l p  to d e f r a y  yield  resource pulp  quantity  from these o v e r l y dense  the c o s t s of r e t u r n i n g such areas  to  production. Although u t i l i z a t i o n of t h i s resource  i n the f o r e s e e a b l e  i s d o u b t f u l , 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 a.  The  include:  s m a l l s i z e o f t h i s m a t e r i a l enhances the p r o s p e c t s  mechanized h a u l i n g and h a n d l i n g (i.e.,  a ' f i b r e combine  1  ( i . e . , bales)  fibres  and  harvesting  However, because the  found i n the b a r k o f d e c i d u o u s s p e c i e s  long  (Crossley,  1938)  a r e absent i n the b a r k o f c o n i f e r o u s s p e c i e s , bark removal  prior  to c o o k i n g  may  complicate  the development o f a  combine' s u i t a b l e f o r use w i t h c o n i f e r o u s  species.  improved methods f o r s e p a r a t i n g bark from c h i p s i s b.  of  such as t h a t r e p o r t e d on i n the  B r i t i s h Columbia Lumberman (1971)). bast  future  U t i l i z a t i o n o f dense stands d u c t i v e s t a t e and  'fibre Research  on  proceeding.  c o u l d r e t u r n l a n d t o a more pro-  t h i s a s p e c t might f a c i l i t a t e n e g o t i a t i o n s f o r  reduced stumpage charges or g r a n t s - i n - a i d of improved management. c.  Present  b o u n d a r i e s between commercial and  might be ered  non-commercial f o r e s t s  r e v i s e d s u b s t a n t i a l l y , because s i t e s p r e s e n t l y  consid-  i n c a p a b l e o f growing t r e e s to merchantable s i z e s might  become  operable.  117  CHAPTER VII  SUMMARY AND SUGGESTIONS FOR FURTHER RESEARCH  The objective of this thesis i s 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 100year-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 aboveground 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 s i t e . The relationships between several crown parameters and radial, cross-sectional area and section volume growth of 100-year-old pine trees were examined in Chapter 4.  lodgepole  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, b a r k - f r e e f u l l b o l e ( 1 . 0 - i n c h t o p ) . were used  i n t h i s study,  warranted.  f u r t h e r development of these r e l a t i o n s h i p s i s  K r a f t pulp y i e l d  ponent and  related  growth r e s u l t s  Because o n l y t e n t r e e s  and  to t r e e s i z e .  q u a l i t y d a t a were o b t a i n e d f o r each comThe  results  i n a s l i g h t l y higher y i e l d  apparent  a f f e c t on p u l p q u a l i t y .  ficantly  related  to the y i e l d  indicated that faster tree  o f p u l p from the f u l l b o l e w i t h  T r e e s i z e and  growth r a t e were not  and q u a l i t y of pulp from the o t h e r  no  signi-  components.  Because the q u a n t i t y of wood f i b r e c o n t a i n e d i n the non-merchantable top ( 4 . 0 - i n c h to 1.0-inch  top) of t r e e s 4.0  to 8.0  i n c h e s i n diameter, which  may  make up a s u b s t a n t i a l p r o p o r t i o n of mature l o d g e p o l e p i n e s t a n d s , i s l a r g e , and because the y i e l d  and  q u a l i t y o f p u l p from t h i s component i s h i g h , immed-  i a t e c o n s i d e r a t i o n s h o u l d be Although  the y i e l d  and  g i v e n to the u t i l i z a t i o n of t h i s  component.  q u a l i t y of p u l p from the root-stump system a r e  i v e l y h i g h , u t i l i z a t i o n of t h i s  component i n the near  relat-  future i s doubtful  because of the t e c h n i c a l problems a s s o c i a t e d w i t h e x t r a c t i n g , c l e a n i n g , t r a n s p o r t i n g , and  p r o c e s s i n g t h i s component.  The  relatively  low y i e l d  and  q u a l i t y of p u l p from branchwood c o u p l e d w i t h the problems o f p r o c e s s i n g t h i s component suggest doubtful.  The  t h a t imminent u t i l i z a t i o n o f l o d g e p o l e p i n e branches i s  y i e l d and  q u a l i t y of p u l p from h i g h l y suppressed  p i n e t r e e s were e s s e n t i a l l y e q u a l to the y i e l d  and  100-year-old  q u a l i t y of pulp  from  mature l o d g e p o l e p i n e . The  r e s u l t s p r e s e n t e d h e r e i n i n no way  e f f e c t s of complete-tree of  attempt  t o examine the  u t i l i z a t i o n on the f o r e s t ecosystem.  Future  study  n u t r i t i o n a l make-up o f the v a r i o u s components o f t r e e s growing under  v a r i o u s f o r e s t c o n d i t i o n s , s i m i l a r to t h a t r e p o r t e d by B o y l e and Ek i s necessary  to e s t a b l i s h  taxes the f o r e s t s i t e . u t i l i z a t i o n on s o i l  the degree to which c o m p l e t e - t r e e  In a d d i t i o n , study of the e f f e c t s o f  (1972),  utilization complete-tree  e r o s i o n , s o i l m o i s t u r e p a t t e r n s and w i l d l i f e management  120  are warranted. Obviously a considerable amount of research must be devoted to the development of methods and equipment before the completetree u t i l i z a t i o n , in whole, w i l l become a viable economic proposition.  121  REFERENCES CITED  Adamovich, L . L . 1970. C e n t r e o f g r a v i t y p o s i t i o n s o f open- and s t a n d grown second growth w e s t e r n c o n i f e r s . Amer. Soc. Agr. Eng., , Pap. No. 70-617. 30 pp. A l e x a n d e r , R. R., D. T a c k l e , and W. G. Dahms. 1967. S i t e indexes f o r l o d g e p o l e p i n e w i t h c o r r e c t i o n s f o r stand d e n s i t y : methodology U. S. Dep. Agr., F o r e s t S e r v . , U. S. F o r e s t Serv. Res. Pap. RM-29. 18 pp. American S o c i e t y f o r T e s t i n g and M a t e r i a l s . 1968. T e n t a t i v e methods o f t e s t f o r s p e c i f i c g r a v i t y o f wood and wood-base m a t e r i a l s , (D 2395-65T). i n 1968 Book o f ASTM Standards. P a r t 16. S t r u c t u r a l Sandwich C o n s t r u c t i o n s ; Wood; A d h e s i v e s . Published by American S o c i e t y f o r T e s t i n g and M a t e r i a l s . P h i l a d e l p h i a . 930 pp. Ando, T.  1965. E s t i m a t i o n o f d r y - m a t t e r and growth a n a l y s i s o f t h e young stand o f Japanese b l a c k p i n e (Pinus t h u n b e r g i i ) . Advanc. F r o n t . P I . S c i . , New D e l h i 10:1-10.  A r t , H. W., and P. L . Marks. 1971. A summary t a b l e o f biomass and n e t annual primary p r o d u c t i o n i n f o r e s t ecosystems o f the w o r l d , i n F o r e s t biomass s t u d i e s . Univ. Maine, L i f e S c i . Agr. Exp. S t a . , M i e s . P u b l . 132, 3-32. Assman, E.  Attiwill,  1961. The p r i n c i p l e s o f f o r e s t y i e l d s t u d y . (English transl a t i o n (1970) by S. H. G a r d i n e r ) . Pergamon P r e s s , T o r o n t o . 506 pp.  P. M. 1966. A method f o r e s t i m a t i n g crown weight i n E u c a l y p t u s , and some i m p l i c a t i o n s o f r e l a t i o n s h i p s between crown weight and stem d i a m e t e r . E c o l . 47_ (5) : 795-804.  B a s k e r v i l l e , G. L . stands. .  1965a. Dry-matter p r o d u c t i o n i n immature balsam f i r F o r e s t S c i . Manogr. 9. 42 pp. 1965b.  Estimation  o f d r y weight o f t r e e components and  total standing crop i n conifer stands. .  1966. Dry-matter production in immature balsam f i r  stands: roots, l e s s e r vegetation, 12 ( 1 ) : 49-53.  and Oak  Ecol. 46 (6): 867-869,  and t o t a l s t a n d .  Forest S c i .  . 1970. T e s t i n g t h e u n i f o r m i t y of v a r i a n c e i n a r i t h m e t i c l o g a r i t h m i c u n i t s o f a Y - v a r i a b l e f o r c l a s s e s o f an X - v a r i a b l e . Ridge Nat. Lab. P u b l . 0RNL-IBP-70-1. 28 pp.  . 1972. Use o f l o g a r i t h m i c r e g r e s s i o n i n t h e e s t i m a t i o n o f p l a n t biomass. Can. J . F o r e s t Res. 2. ( 1 ) : 49-53.  122  B a z i l e v i c , N. I . , and L. E. R o d i n . 1966. The b i o l o g i c a l c y c l e o f n i t r o g e n and ash elements i n p l a n t communities o f the t r o p i c a l and s u b t r o p i c a l zones. F o r e s t . A b s t r . L e a d i n g A r t . Ser. No. 38, F o r e s t . A b s t r . 27 ( 3 ) : 357-368. Bella,  1. E.  1968. E s t i m a t i n g a e r i a l component w e i g h t s of young aspen trees. Can. Dep. F o r e s t . R u r a l Develop., F o r e s t . Br., F o r e s t Res. Lab., Inform. Rep. MS-X-12. 36 pp. 1970. S i m u l a t i o n of growth, y i e l d , and management o f aspen, Fac. F o r e s t . , U n i v . B r i t i s h Columbia, Ph.D. T h e s i s . 190 pp. and J . M. aspen s t a n d WR-437.  Jarvis. 1967. H i g h t o t a l p r o d u c t i v i t y of a young i n Manitoba. P u l p Pap. Mag. Can. 68_ (10):WR-432,  Bormann, F. H., G. E. L i k e n s , D. W. F i s h e r , and R. S. P i e r c e . 1967. N u t r i e n t l o s s a c c e l e r a t e d by c l e a r c u t t i n g a f o r e s t e c o s y s t e m . i n Symposium on p r i m a r y p r o d u c t i v i t y and m i n e r a l c y c l i n g i n n a t u r a l ecosystems. U n i v . Maine P r e s s , Orono, Maine. 187-196. Boyle,  J . R., and A. R. Ek. 1972. An e v a l u a t i o n of some e f f e c t s o f b o l e and branch pulpwood h a r v e s t i n g on s i t e m a c r o n u t r i e n t s . Can. J . F o r e s t Res. 2 ( 4 ) : 407-412.  B r i t i s h Columbia Lumberman. 1971. New system d e l i v e r s c h i p s to m i l l a t h a l f the c o s t . B r i t i s h Columbia Lumberman 55 ( 7 ) : 47-48. B u r g e r , H.  1929. H o l z , B l a t t menge und Zuwachs. I . D i e Weymouthsfore. M i t t e i l . Schweiz. C e n t r a l a n s t . F o r s t l . Versuchsw. 15 ( Z ) : 243292.  Chase, A. J . , F. Hyland, and H. E. Young. 1971. Puckerbrush pulping s t u d i e s . Univ. Maine, L i f e S c i . Agr. Exp. S t a . , Tech. B u l l . 49. 64 pp. C h i d e s t e r , G. H., M. W. Bray, and C. E. C u r r a n . 1939. Growth r a t e and p o s i t i o n o f wood i n t r e e as f a c t o r s i n f l u e n c i n g k r a f t and s u l f i t e pulps from j a c k p i n e . J . F o r e s t . 3_7 ( 9 ) : 680-683. C o l e , W.  D.,  S. P. G e s s e l , and S. F. D i c e . 1967. D i s t r i b u t i o n and c y c l i n g o f n i t r o g e n , phosphorous, potassium and c a l c i u m i n a secondgrowth D o u g l a s - f i r ecosystem. In Symposium on primary p r o d u c t i v i t y and m i n e r a l c y c l i n g i n n a t u r a l ecosystems. U n i v . Maine P r e s s , Orono, Maine. 197-232.  C r o s s l e y , T. L. 1938. Paper from f r u i t t r e e p r u n i n g s and P u l p Pap. Mag. Can. 39 ( 8 ) : 568-570. Crow, T.  R.  D e n g l e r , A.  forest slash.  1971. E s t i m a t i o n of biomass i n an even-aged s t a n d - r e g r e s s i o n and "mean t r e e " t e c h n i q u e s , i n F o r e s t biomass s t u d i e s . Univ. Maine, L i f e S c i . Agr. Exp. S t a . , M i s c . P u b l . 132, 35-48. 1937. Krownengrosse, Nadelmenge und Z u w a c h s l e i s t u n g A i t k i e f e r n . Z e i t s c h r . F o r s t Jgdw. 69 ( 7 ) : 321-336.  van  123  Duvigneaud, P., and S. Denaeyer-DeSmet. 1967. Biomass, p r o d u c t i v i t y and m i n e r a l c y c l i n g i n deciduous f o r e s t s i n B e l g i u m . i n Symposium on primary p r o d u c t i v i t y and m i n e r a l c y c l i n g i n n a t u r a l ecosystems. U n i v . Maine P r e s s , Orono, Maine. 167-186. Dyer, R.  F.  1967. F r e s h and d r y w e i g h t , n u t r i e n t elements and p u l p i n g c h a r a c t e r i s t i c s o f n o r t h e r n w h i t e cedar Thuja o c c i d e n t a l i s . Maine Agr. Exp. S t a . , Tech. B u l l . 27. 40 pp. A. J . Chase, and H. E. Young. 1968. P u l p from p r e s e n t l y non-commercial woody p e r e n n i a l s . P u l p Pap. Mag. Can. 69_ ( 1 ) : 57-62.  E l l e n b e r g , H. 1971. E c o l o g i c a l s t u d i e s 2. Integrated experimental ecology Methods and r e s u l t s o f ecosystem r e s e a r c h i n the German S o i l i n g Project. (H. E l l e n b e r g , E d i t o r ) . S p r i n g e r - V e r t a g , New York 214 pp. F a r r a r . J . L. 1961. L o n g i t u d i n a l v a r i a t i o n i n the t h i c k n e s s of the ring. F o r e s t . Chron. 3_7 ( 4 ) : 323-330. H a l l , G.  Hatton,  S.  annual  1965. Wood increment and crown d i s t r i b u t i o n r e l a t i o n s h i p s i n r e d p i n e : F o r e s t S c i . 11 ( 4 ) : 438-448.  J . V. 1972. Res. S c i . ( P u l p i n g ) , Western F o r e s t P r o d u c t s L a b o r a t o r y , Vancouver. P e r s o n a l communication - u n p u b l i s h e d data. . and J . L. Keays. 1970. R e l a t i o n s h i p between p u l p y i e l d and permanganate number f o r k r a f t p u l p s : 1 western hemlock, white spruce and l o d g e p o l e p i n e . P u l p Pap. Mag. Can. _71 (11-12) : T259-T268. and J . L. Keays. 1971. Laboratory p r e c i s i o n d i g e s t e r studies. 1. l a c k o f i n t e r a c t i o n i n k r a f t p u l p i n g mixtures western hemlock t r e e components. Svensk P a p p e r s t i d . 74 ( 5 ) : 128-131.  of  and M. Samkova. 1972. R e l a t i o n s h i p between b u l k and handsheet p h y s i c a l p r o p e r t i e s f o r k r a f t p u l p s . 1. Tree components o f P i c e a g l a u c a . T a p p i 55 ( 1 ) : 93-96. Heger, L.  1965. Morphogenesis o f stems o f Douglas f i r (Pseudotsuga m e n z i e s i i (Mirb.) F r a n c o ) . Fac. F o r e s t . , Univ. o f B r i t i s h Columbia, Ph.D. T h e s i s . 176 pp. p l u s a p p e n d i c e s .  Hegyi,  1972. Dry matter d i s t r i b u t i o n i n j a c k p i n e stands Ontario. F o r e s t . Chron. 48 ( 4 ) : 193-197.  F.  Honer, T. G.  i n northern  1971. Weight r e l a t i o n s h i p s i n open- and f o r e s t - g r o w n balsam f i r t r e e s , i n F o r e s t biomass s t u d i e s . Univ. Maine, L i f e S c i . Agr. Exp. S t a . , M i s c . P u b l . 132, 65-78.  124  H o r t o n , K.  W. 1956. The ecology o f l o d g e p o l e p i n e i n A l b e r t a and i t s role i n forest succession. Can. Dep. N. A f f . Nat. Resources, F o r e s t . Br., F o r e s t Res. D i v . , Tech. Note 45. 29 pp. . 1958. R o o t i n g h a b i t s o f l o d g e p o l e p i n e . Can. Dep. N. A f f . Nat. Resources, F o r e s t . Br., F o r e s t Res. D i v . , Tech. Note 67 26 pp.  Hunt, K.  •  1971. A comparison o f k r a f t p u l p i n g o f sound and dwarf m i s t l e t o e i n f e c t e d western hemlock wood. Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . S e r v . , F o r e s t Prod. Lab., Inform. Rep. VP-X-78. 7 pp. .  and J . L. Keays. 1970. P r e l i m i n a r y r e p o r t on the p u l p i n g o f l o d g e p o l e p i n e i n f e c t e d w i t h a t r o p e l l i s canker. Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . Serv., F o r e s t Prod. Lab., I n t e r n . Rep. VP-56. 17 pp.  J o h n s t o n e , W. D. 1967. A n a l y s i s of biomass, biomass sampling methods, and weight s c a l i n g o f l o d g e p o l e p i n e . Fac. F o r e s t . , Univ. B r i t i s h Columbia, M.F. T h e s i s . 153 pp. '  . 1970a. Some v a r i a t i o n s i n s p e c i f i c g r a v i t y and m o i s t u r e c o n t e n t of 1 0 0 - y e a r - o l d l o d g e p o l e p i n e t r e e s . Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . Serv., F o r e s t . Res. Lab., Inform. Rep. A-X-29. 19 pp. '  . 1970b. Component d r y - w e i g h t s o f 1 0 0 - y e a r - o l d l o d g e p o l e pine t r e e s . Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . Serv., F o r e s t Res. Lab., Inform. Rep. A-X-31. 14 pp. . 1971. T o t a l s t a n d i n g crop and t r e e component d i s t r i b u t i o n s i n t h r e e s t a n d s o f 1 0 0 - y e a r - o l d l o d g e p o l e p i n e , i n F o r e s t biomass studies. U n i v . Maine, L i f e S c i . Agr. Exp. S t a . , M i s c . P u b l . 132, 81-89.  • .  '  1972. Unpublished  F o r . O f f . , N. data.  F o r e s t Res.  Lab.,  Edmonton.  Keays, J . L.  1968. Whole-tree u t i l i z a t i o n s t u d i e s : s e l e c t i o n o f t r e e components f o r p u l p i n g r e s e a r c h . Can. Dep. F o r e s t . R u r a l Develop., F o r e s t . Br., F o r e s t Prod. Lab., Inform. Rep. VP-X-35. 31 pp.  __.  1970. World developments i n i n c r e a s e d f o r e s t r e s o u r c e s f o r the pulp and paper i n d u s t r y . Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . Serv., F o r e s t P r o d . Lab., Inform. Rep. VP-X-64. 64 pp.  .  . 1971a. C o m p l e t e - t r e e u t i l i z a t i o n . Resume of a l i t e r a t u r e review, i n F o r e s t biomass s t u d i e s . Univ. Maine, L i f e S c i . Agr. Exp. S t a . , M i s c . P u b l . 132, 93-102.  125  1971b. C o m p l e t e - t r e e u t i l i z a t i o n . An a n a l y s i s o f the literature. P a r t I : Unmerchantable top of b o l e . Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . S e r v . , F o r e s t P r o d . Lab., Inform. Rep. VP-X-69. 98 pp. . 1971c. Complete-tree u t i l i z a t i o n . An a n a l y s i s o f the literature. P a r t I I : F o l i a g e . Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . S e r v . , F o r e s t P r o d . Lab., Inform. Rep. VP-X-70. 94 pp. . 1971d. C o m p l e t e - t r e e u t i l i z a t i o n . An a n a l y s i s o f the l i t e r a t u r e . P a r t I I I : B r a n c h e s . Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . S e r v . , F o r e s t P r o d . L a b . , Inform. Rep. VP-X-71. 67 pp. . 1971e. C o m p l e t e - t r e e u t i l i z a t i o n . An a n a l y s i s o f the literature. P a r t IV: Crown and s l a s h . Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . S e r v . , F o r e s t P r o d . L a b . , Inform. Rep. VP-X-77. 79 pp. . 1971f. Complete-tree u t i l i z a t i o n . An a n a l y s i s o f the literature. P a r t V: Stump, r o o t , and stump-root system. Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . S e r v . , F o r e s t P r o d . Lab., I n f o r m . Rep. VP-X-79. 62 pp. . 1972. Res. S c i . ( P u l p i n g ) , Western F o r e s t P r o d u c t s L a b o r a t o r y . Vancouver, P e r s o n a l communication - u n p u b l i s h e d data. . and J . M. B a g l e y . 1970. pulping studies. T a p p i 53  D i g e s t e r assembly ( 1 0 ) : 1935-1940.  for precision  . and J . V. H a t t o n . 1971a. C o m p l e t e - t r e e u t i l i z a t i o n s t u d i e s . I. Y i e l d and q u a l i t y o f k r a f t p u l p from the components o f Tsuga h e t e r o p h y l l a . T a p p i 54 ( 1 ) : 99-104. . and J . V. H a t t o n . 1971b. Complete-tree u t i l i z a t i o n s t u d i e s . II. Y i e l d and q u a l i t y o f k r a f t pulp from the components of P i c e a g l a u c a . T a p p i 54 ( 1 0 ) : 1721-1724. . and J . V. H a t t o n , and R. R. C o r t e z . 1969. The p r e c i s i o n o f l a b o r a t o r y k r a f t p u l p - y i e l d d e t e r m i n a t i o n s . T a p p i _52 ( 5 ) : 904912. Keen. R. E.  1963. Weights and c e n t r e s o f g r a v i t y i n v o l v e d i n h a n d l i n g pulpwood t r e e s . P u l p Pap. Res. I n s t . , Can., Woodlands Res. Index 147. 93 pp.  K i i l , A. D.  1965. S l a s h weight and s i z e d i s t r i b u t i o n o f w h i t e . s p r u c e and l o d g e p o l e p i n e . F o r e s t Chron. 4_1 (4) : 432-437.  126  1968. Changes i n the p h y s i c a l c h a r a c t e r i s t i c s and m o i s t u r e c o n t e n t o f p i n e and s p r u c e - f i r s l a s h d u r i n g the f i r s t f i v e y e a r s a f t e r l o g g i n g . Can. Dep. F o r e s t . R u r a l Develop., F o r e s t . B r . , I n t e r n . Rep. A-14. 40 pp. K i r a , T.,  and T. S h i d e i . 1967. P r i m a r y p r o d u c t i o n and t u r n o v e r o f o r g a n i c m a t t e r i n d i f f e r e n t f o r e s t ecosystems o f the w e s t e r n P a c i f i c . Jap. J . E c o l . 17 ( 2 ) : 70-87.  K i r b y , C. L.  1968. S i t e i n d e x and h e i g h t growth p r e d i c t i o n s f o r dominant and codominant w h i t e s p r u c e and l o d g e p o l e p i n e i n A l b e r t a . Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . S e r v . , F o r e s t Res. Lab., (mimeo), 10 pp.  Kittredge, J . 1944. E s t i m a t i o n o f the amount o f f o l i a g e of t r e e s stands. J . F o r e s t . 42 (12): 905-912. Kozak, A.,  and J . H. G. Smith. 1965. A comprehensive and f l e x i b l e m u l t i p l e r e g r e s s i o n program f o r e l e c t r o n i c computing. Forest. Chron. 41 ( 4 ) : 438-443.  Kramer, P.J. , and T. T. K o z l o w s k i . H i l l Book Co. I n c . , New Kurucz, J .  L e e , Y.  and  1960. York,  Physiology of trees. 642 pp..  McGraw-  1969. Component weights o f Douglas f i r , w e s t e r n hemlock, and w e s t e r n r e d cedar biomass f o r s i m u l a t i o n o f amount and d i s t r i bution of forest fuels. Fac. F o r e s t . , U n i v . B r i t i s h Columbia, M.F. T h e s i s . 116 pp.  1967. Stand models f o r l o d g e p o l e p i n e and l i m i t s to t h e i r application. Fac. F o r e s t . , Univ. B r i t i s h Columbia, Ph.D. Thesis. 332 pp.  MacDonald, R. G. 1969. McGraw-Hill  P u l p and Paper Manufacture. Book Co. I n c . , New York. 769  V o l . I. 2nd pp.  Edition  Madgwick, H.A.I. 1970. Biomass and p r o d u c t i v i t y models o f f o r e s t canopies, in Ecological studies. 1. A n a l y s i s o f temperate f o r e s t ecosystems. (D. E. R e i c h l e , E d i t o r ) . S p r i n g e r - V e r l a g , New York: 47-54. Marchenko, A. I . 1967. P e c u l i a r i t i e s of biogeochemical processes i n spruce f o r e s t s and f i l l e d a r e a s , i n Symposium on p r i m a r y p r o d u c t i v i t y and m i n e r a l c y c l i n g i n n a t u r a l ecosystems. Univ. Maine P r e s s , Orono, Maine: 139-150. M a r : M o l l e r , C. 1947. The e f f e c t o f t h i n n i n g , age, and s i t e on f o l i a g e , i n c r e m e n t , and l o s s o f dry m a t t e r . J . F o r e s t . 4-5 ( 6 ) : 393-  404. Mason, D.  I.  1915. The l i f e h i s t o r y o f l o d g e p o l e p i n e i n the Rocky Mountains. U. S. Dep. Agr., F o r e s t Serv., B u l l . 154. 35  pp.  127  McGreevy, M.  G. 1972. A s i m u l a t i o n o f t h e t r e e components o f the f o r e s t f u e l complex t o a i d i n p l a n n i n g f o r f i r e c o n t r o l use. F a c . F o r e s t . , U n i v . B r i t i s h Columbia, M.F. T h e s i s . 121 pp.  Meyer, H. A.  1944. A c o r r e c t i o n f o r a s y s t e m a t i c e r r o r o c c u r r i n g i n t h e a p p l i c a t i o n o f t h e l o g a r i t h m i c volume e q u a t i o n . Penn. S t a t e F o r e s t . Sch., Res. Pap. No. 7. 3 pp.  Moir , W. H., and R. F r a n c i s . 1972. F o l i a g e biomass and s u r f a c e a r e a i n three Pinus c o n t o r t a p l o t s i n Colorado. Forest. S c i . 18 ( 1 ) : 41-45. Monsi, M.  1968. M a t h e m a t i c a l models o f p l a n t communities, i n F u n c t i o n i n g o f t e r r e s t r i a l ecosystems a t t h e primary p r o d u c t i o n l e v e l . P r o c . Copenhagen Symp. ( F . E. E c k a r d t , E d i t o r ) , UNESCO, P a r i s : 131-150.  Munro, D. D.  1970. The u s e f u l n e s s o f form measures i n t h e e s t i m a t i o n o f volume and t a p e r o f the commercial t r e e s p e c i e s o f B r i t i s h Columbia. Sec. 25, I.U.F.R.O., B i r m e n s d o r f , S w i t z e r l a n d , (mimeo), 13 pp.  Muraro, S. J . 1966. Lodgepole p i n e l o g g i n g s l a s h . P u b l . No. 1153. 14 pp.  Can. Dep. F o r e s t . ,  N i c h i p o r o v i c h , A. A. 1967. P h o t o s y n t h e s i s o f p r o d u c t i v e systems. (A. A. N i c h i p o r o v i c h , E d i t o r ) , I s r a e l Program f o r S c i e n t i f i c Translations Ltd., Jerusalem. 182 pp, O l s o n , D. F. J r . 1 1971. Sampling l e a f biomass i n even-aged stands o f y e l l o w p o p l a r ( L i r i o d e n d r o n t u l i p i f e r a L . ) . i n F o r e s t biomass studies. U n i v . Maine, L i f e S c i . A g r . Exp. S t a . , M i s c . P u b l . 132. 115-122. Onaka, F.  1950a. The l o n g i t u d i n a l d i s t r i b u t i o n o f r a d i a l increment i n trees. Kyoto U n i v . F o r e s t . B u l l . 18: 1-53.  1950b. The effects of such treatments as defoliation, d i s budding, girdling, and screening of light on growth and especially radial growth of evergreen conifers. Kyoto Univ. Forest. B u l l . 18: 55-95. Orman, H. R., and G. M. W i l l . 1960. The n u t r i e n t c o n t e n t o f P i n u s r a d i a t a t r e e s . New Z e a l . J . S c i . 3 ( 3 ) : 510-522. Osborn, J . E. 1968. I n f l u e n c e o f s t o c k i n g and d e n s i t y upon growth and y i e l d o f t r e e s and stands o f c o a s t a l w e s t e r n hemlock. Fac. F o r e s t . , Univ. B r i t i s h Columbia, Ph.D. T h e s i s . 396 pp. O v i n g t o n , J . D. 1956. The form, w e i g h t s and p r o d u c t i v i t y o f t r e e grown i n c l o s e s t a n d s . New P h y t o l . _55 ( 2 ) : 289-304.  species  128  . 1962. Q u a n t i t a t i v e ecology and the woodland ecosystem concept. Advances E c o l . Res. 1_: 103-192. . and H. A. I . Madgwick. 1959. D i s t r i b u t i o n o f o r g a n i c m a t t e r and p l a n t n u t r i e n t s i n a p l a n t a t i o n o f Scots p i n e . F o r e s t S c i . _5 (4) : 344-355. Paille,  G.  1970. D e s c r i p t i o n and p r e d i c t i o n o f m o r t a l i t y i n some c o a s t a l Douglas f i r s t a n d s . F a c . F o r e s t . , Univ. B r i t i s h Columbia, Ph.D. T h e s i s . 300 pp.  P a r k e r , M. L . , and W. E. S. Henoch. 1971. The use o f Engelmann latewood d e n s i t y f o r d e n d r o c h r o n o l o g i c a l p u r p o s e s . Can. J . F o r e s t Res. 1. ( 2 ) : 90-98. Peterson,  spruce  E. B?' 1970. Methods f o r e s t i m a t i n g s t a n d i n g crop i n Populus f o r e s t s o f A l b e r t a . Can. Dep. F i s h . F o r e s t . , Can. F o r e s t . Serv., F o r e s t Res. Lab., I n t e r n . Rep. A-29. 22 pp. 1972. Dep. E n v i r o n . , N. F o r e s t Res. Lab., Edmonton, A l t a . Personal correspondence. , Y. H. Chan, and J . B. Cragg. 1970. Aboveground s t a n d i n g c r o p , l e a f a r e a , and c a l o r i c v a l u e i n an aspen c l o n e near C a l g a r y , A l b e r t a . Can. J . B o t . 48 ( 7 ) : 1459-1469.  P o l l a r d , D. F. W. stands P o s t , L. J .  1972. Above-ground d r y matter p r o d u c t i o n i n t h r e e o f t r e m b l i n g aspen. Can. J . F o r e s t . Res. 2^ ( 1 ) : 27-33.  1970. Dry-matter p r o d u c t i o n o f mountain maple and balsam f i r i n n o r t h w e s t e r n New Brunswick. E c o l . J51_ (3) : 548-550.  P u l p and Paper Research I n s t i t u t e o f Canada. 1962. The l a b o r a t o r y p r o c e s s i n g o f p u l p (PFI m i l l method). PPRIC Standard T e s t P r o c e d u r e No. PB-6. 5 pp. Rennie, P. J . 1955. The uptake o f n u t r i e n t s by mature f o r e s t P l a n t S o i l 1_ (1) : 49-95.  growth.  Rodin, L . E . , and N. I . B a z i l e v i c h . 1965. P r o d u c t i o n and m i n e r a l c y c l i n g i n t e r r e s t i a l vegetation. ( E n g l i s h t r a n s l a t i o n (1967) by G. E . Hogg). O l i v e r and Boyd L t d . , London. 288 pp. , and N. I . B a z i l e v i c h . The b i o l o g i c a l p r o d u c t i v i t y o f the main v e g e t a t i o n types i n n o r t h e r n hemisphere o f the o l d w o r l d . F o r e s t . A b s t r . L e a d i n g A r t . S e r . No. 38, F o r e s t . A b s t r . 27 ( 3 ) : 369-372. , and N. I . B a z i l e v i c h . 1968. World d i s t r i b u t i o n o f p l a n t biomass. I n F u n c t i o n i n g o f t e r r e s t r i a l ecosystems a t the primary production l e v e l . P r o c . Copenhagen Symp. ( F . E. E c k a r d t , E d i t o r ) , UNESCO, P a r i s : 45-52.  129  Rowe, J . S.  Sastry,  1959. F o r e s t r e g i o n s o f Canada. Resources, F o r e s t . B r . , B u l l . 123.  Can. Dep. 71 pp.  C. B. R. , A. Kozak, and R. W. Wellwood. 1972. f o r the e v a l u a t i o n o f wood from f e r t i l i z e d F o r e s t Res. 2 ( 4 ) : 417-426.  Satoo, T.  N. A f f .  Nat.  A new approach trees. Can. J .  1967. P r i m a r y p r o d u c t i o n r e l a t i o n s i n woodlands of Pinus d e n s i f l o r a . i n Symposium on p r i m a r y p r o d u c t i v i t y and m i n e r a l c y c l i n g i n n a t u r a l ecosystems. Univ. Maine P r e s s . Orono, Maine: 52-80.  Shea, S. R.,  Shinozaki,  and K. A. Armson. 1972. Stem a n a l y s i s o f j a c k p i n e (Pinus b a n k s i a n a , Lamb.): t e c h n i q u e s and c o n c e p t s . Can. J . F o r e s t Res. 2 ( 4 ) : 392-406.  K., K. Yoda, K. Hozumi, and T. K i r a . 1964. A quantitative a n a l y s i s of p l a n t form - the p i p e model t h e o r y . I. Basic analyses. Jap. J . E c o l . 14 ( 9 ) : 97-105.  Smith, J . H.  G. 1971. Bases f o r sampling and s i m u l a t i o n i n s t u d i e s of t r e e and s t a n d weight, i n F o r e s t biomass s t u d i e s . Univ. Maine. L i f e S c i . Agr. Exp. S t a . , M i s c . P u b l . 132, 139-149. . 1972. P r o f . , Fac. F o r e s t . , Univ. B r i t i s h Columbia, Vancouver, B.C. P e r s o n a l communication - u n p u b l i s h e d data. , and A. Kozak. 1967. T h i c k n e s s and p e r c e n t a g e o f bark o f the commercial t r e e s o f B r i t i s h Columbia. Fac. F o r e s t . , Univ. B r i t i s h Columbia, (mimeo). 27 pp.  S m i t h e r s , L. A. 1957. T h i n n i n g i n l o d g e p o l e p i n e stands i n A l b e r t a . Can. Dep. N. A f f . Nat. Resources, F o r e s t . B r . , F o r e s t Res. D i v . , Tech. Note 52. 26 pp. . 1961. B u l l . 127. S p r o u l l , R.  Lodgepole pine 153 pp.  i n A l b e r t a . Can.  C., R. B. P a r k e r , and W. L. B e l v i n . 1957. i n g . F o r e s t Prod. J . 1_ ( 4 ) : 131-134.  Dep.  Forest.,  Whole t r e e  harvest-  Steinbeck,  K., and J . T. May. 1971. P r o d u c t i v i t y o f v e r y young P l a n t a n u s o c c i d e n t a l i s L. p l a n t i n g s grown at v a r i o u s s p a c i n g s . i n F o r e s t biomass s t u d i e s . U n i v . Maine, L i f e S c i . Agr. Exp. S t a . , M i s c . P u b l . 132, 153-162.  S t i e l l , W.  M. 1966. Can. Dep.  Szabo, T.  Red p i n e crown development i n r e l a t i o n to F o r e s t . , P u b l . No. 1145. 44 pp.  spacing.  and J . L. Keays. 1973. P u l p i n g o f spruce and p i n e tops from Alberta. Can. Dep. E n v i r o n . , Can. F o r e s t . Serv., F o r e s t P r o d . Lab., ( i n p r e p a r a t i o n ) . 6 pp.  130  1966. Some dimensions on the l e a f biomass of f o r e s t stands and t r e e s . B u l l . Govt. F o r e s t Exp. S t a . , No. 184., Tokyo,  T a d a k i , Y.  135-159. _, N. Ogata, and T. T a k a g i . 1962. S t u d i e s on p r o d u c t i o n s t r u c t u r e of forest ( I I I ) . E s t i m a t i o n o f s t a n d i n g crop and some a n a l y s e s on p r o d u c t i v i t y of young stands o f G a s t a n o p s i s c u s p i d a t a . J . J a p . F o r e s t . Soc. 44 ( 1 2 ) : 350-359. Wassink, E. C. 1968. L i g h t energy c o n v e r s i o n i n p h o t o s y n t h e s i s and growth of p l a n t s , i n F u n c t i o n i n g of t e r r e s t r i a l ecosystems a t the primary p r o d u c t i o n l e v e l . P r o c . Copenhagen Symp. (F. E. E c k a r d t , E d i t o r ) , UNESCO, P a r i s : 53-66. weetman, G.  F., and R. H a r l a n d . 1964. F o l i a g e and wood p r o d u c t i o n i n unthinned b l a c k s p r u c e i n n o r t h e r n Quebec. F o r e s t S c i . 10 ( 1 ) : 80-88.  White, E. H., W. L. P r i t c h e t t , and W. K. Robertson. 1971. Slash pine root biomass and n u t r i e n t c o n c e n t r a t i o n s , in F o r e s t biomass s t u d i e s . Univ. Maine, L i f e S c i . Agr. Exp. S t a . , M i s c . P u b l . 132, 165-176. Whittaker,  Will,  R. H. 1966. F o r e s t dimensions and p r o d u c t i o n i n the Smoky Mountains. E c o l . 4_7 ( 1 ) : 103-121.  G. M.  Great  1964. Dry m a t t e r p r o d u c t i o n and n u t r i e n t uptake by Pinus r a d i a t a i n Nexvf Z e a l a n d . Comm. F o r e s t . Rev. 43 ( 1 ) : 57-70.  Woodwell, G. M., and R. H. W h i t t a k e r . 1967. P r i m a r y p r o d u c t i o n and the c a t i o n budget o f the Brookhaven F o r e s t , in Symposium on primary p r o d u c t i v i t y and m i n e r a l c y c l i n g i n n a t u r a l ecosystems. Univ. Maine P r e s s , Orono, Maine: 151-166. Young, H.  E.  1964. The complete t r e e concept - A c h a l l e n g e and an opportunity. P r o c . Soc. Amer. F o r e s t . , Denver, C o l o . : 231-233.  . 1965a. Pound w i s e and penny f o o l i s h . Annu. Meeting, New York, (mimeo), 9 pp.  Amer. Pulpwood  . 1965b. Is t r e e growth of ten cords per a c r e per attainable? Maine Farm Res. 13 ( 3 ) : 1-7.  Assoc.  year  . 1966. World f o r e s t r y based on the complete t r e e concept. P r o c . S i x t h World F o r e s t . Congr., Madrid, S p a i n : 2835-2839. . 1967. The l o s t c o r d . A r e a IV S e c t . , S. Pulpwood Conserv. Assoc., Nag's Head, N.C., (mimeo). 11 pp. 1968. C h a l l e n g e o f complete t r e e u t i l i z a t i o n . Prod. J . 18 ( 4 ) : 83-86.  Forest  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 i n Maine. Maine Agr. Exp. Sta., Tech. B u l l . 12. 76 pp. , and A. J . Chase. 1965. Fiber weight and pulping characteristics of the logging residue of seven tree species in Maine. Maine Agr. Exp. Sta., Tech. B u l l . 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. B u l l . 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 i n 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 i n biological data. Bio. Sci. IB (12) : 1118-1120.  132  APPENDIX I .  L e s s e r v e g e t a t i o n p r e s e n t i n Stands  A s t e r conspicuus  1 and  Lindl.  Cornus c a n a d e n s i s  2.  Showy a s t e r  L.  Bunchberry  Cornus s t o l o n i f e r a Michx.  Red  Elymus i n n o v a t u s B e a l .  Hairy wild  E p i l o b i u m a n g u s t i f o l i u m L.  Fireweed  Equisetum  Scouring rush  Galium  hyemale L.  l a b r a d o r i c u m Wieg.  rye  Bedstraw  L a t h y r u s o c h r o l e u v u s Hook.  Pea v i n e  L i n n a e a b o r e a l i s L. v a r . americana  ( F o r b e s ) Rehd.  L o n i c e r a d i o i c a L. v a r . g l a u c e n s c e n s P e t a s i t e s palmatus ( A i t . ) A. Populus  o s i e r dogwood  Gray  t r e m u l o i d e s Michx.  (Rydb.) B u t t e r s  Twin-flower Twining  honeysuckle  Palmate-leaved Trembling  coltsfoot  aspen  P y r o l a a s a r i f o l i a Michx.  Common p i n k w i n t e r g r e e n  Rosa a c i c u l a r i s  Prickly  Lindl.  Smilicina s t e l l a t a  (L.) Desf.  S t r e p t o p u s a m p l e x i f o l i u s (L.) T h a l i c t r u m o c c i d e n t a l e A. Viburnum t r i l o b u m Marsh. V i o l a r u g u l o s a Greene  Gray  rose  Star-flowered Seal DC.  Solomon's-  Twisted-stalk Western meadowrue High-bush c r a n b e r r y Western Canada v i o l e t  133  Appendix 1-1.  Appendix H . - 2 . 0 R Y ABOVE-GROUND WE1GHT-DBH ALLOMETRY  DRY TOTAL TREE WEK5HT-DBH ALLOMETRY OF  MATURE WHITE SPRUCE.  OF  MATURE WHITE SPRUCE.  O«NI«,.I  MHI*.I  Appendix B - 3 . DRY STEM WEIGHT-BASAL AREA RELATIONSHIP  Appendix 1 - 4  . DRY MERCHANTABLE STEM WEIGHT-BASAL AREA RELATIONSHIP OF MATURE WHITE  OF MATURE WHITE SPRUCE.  SPRUCE.  •*  *  A I M  -  i  e r a l ATM [«lt]  t  *  IX  M  •  •»  .«  .4  TrM  Jk  total A/*o(iQ.ft.)  10  IJ  u  134  A p p e n d i x B-5.  DRV  NEEDIE  OF M A T U R E  WEIGHT-DBH WHITE  /  Appendix 1 - 6 . DRY  ALLOMETRY  OF  SPRUCE.  / I  "0.9 i  V.*  \ 08H  ALLOMETRY  SPRUCE.  •  /  n - IB  M.2 lb, [24  WEIGHT-DBH  WHITE  • /  IW*2.030 Log^O- 0.163 1 t  BRANCH MATURE  /  8%]  (MV)  A p p e n d i x JT-7.  DRY  ROOT-STUMP  RELATIONSHIP  OF  WEIGHT-BASAL  MATURE  WHITE  AREA  SPRUCE.  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.)  097  n • 18 '» lb. [4141b]  .'•or: 5  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.  dbhob • 6.3 i«. Wight • 34.B fl  . JO-  - 30  S'  3 10 15 5-yoor rodiol Dry woigta of growth Lmn) loliogo (IbJ  Dry woighl of  growth (mm)(Oliogo Hb)  TREE  3  TREE  «bhob- 3.] i„. hoight • 43.3 (i.  dbhab • 3.3 in. •night • 34.3 ft.  3 K) K> 3 3-jroor radial Dry woighl of growth \mm)•ollogo lib}  4  3-yoor radial Ory woight of growth 1»n) loliogo (IbJ  137  TREE'6  TREE 5  tfbhob • 7.0 «. bolglit • oJ.« ft.  • K.Sr>. hoigM • 67.T f«.  4bkob  Eh  5 it) S 20 25 30 35 40 Dry woight of fotiog* fIbJ  TREE  10 J 5 to 13 3-yoor rao*iol Dry w«ig>t of growth (mm) foliogo (IW  7  TREE  ObHob • 7.0 in. Wight - 34.0 h.  J-yOOr rotf%ol  Dry w«.gM of foliogo (IbJ  8  3.8 io. 39.3 ft.  3-yoor  rodiol  ffrowtn (mm)  Dry woighl of foliogo ClbJ  138  JO  40  2 X  30 JO-  »•  0  3-yoor  5-yoor rodiol Dry woight ef foliogo (lb) growth (mm)  Dry Might of radio) foliogo (IbJ  TREE II  TREE 12  dbhob < W . 2 io. hoight ' 7J.J ft.  dbhob • 7.4 ir  70  bight • tl}  to  JO  3-  Jo  1  10 to  »  J  Dry woight Of  3-yoor rodiol  foliogo (lb.)  growth ( » • )  S » U 30 2J Dry woight of foliogo (lb)  139  TREE 14  TREE 13 dbhob • 11.* h. MgM • 77.4 ft  •boob  • 7.2  bight  • «0.7  ov K  i  »  ~3  5  3-ytrar rodiol  »  3  Dty woight  Srowril (mm)  55 5  3  of foliogo  Ob)  5  3 IJ  -  10  3  3- poor todiot  5 Dry  U JO  K  wMgb* of foiiogo (lb)  fjrowrb(M)  TREE 15 dbhob •  TREE 16  7 3 io.  hoight • 39.1 l l .  HZ  n  i  3-yoor rodiol Kfmak)  J  10 13  Dry woighl of foliogo (lb)  w 3-yoor  1 rodiol  growth (oia)  3 Dry  10 13 woighl  foliogo  of  Ob)  dbhob  .  7.J i « .  hoight  •  57J  Jl.  140  TREE  TREE  17  18  .bkob : 4.8 in. bight • 47 2 It.  dbhob • S3 r». k.igr» • 61.7 It. •0  50  40  s  o  I »•  10  30 100  70  » 5 3 n 5-ywor rodiol Dry w.ight growth (mnu of foliog. (lb)  5 3 10 13 rodiol Dry weight of growth iiorn) foliog. (lb) K  5-yMr  TREE 19  TREE 20  dbhob • 8.8 in. hoight • 69.2 ft.  dbhob • 4.7 in. hoight - 36.2 ft.  30402 I *  2  JO-  10-  3 10 13 20 13 Dry w.ight of foliogo (lb)  

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
http://iiif.library.ubc.ca/presentation/dsp.831.1-0101020/manifest

Comment

Related Items