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Weight-length relationships of coniferous wood tracheid skeletons 1971

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WEIGHT-LENGTH RELATIONSHIPS OF CONIFEROUS WOOD TRACHEID SKELETONS by CHERLA BHASKARA RAMA SASTRY M. S c . ( B o t a n y ) ' A n d h r a U n i v e r s i t y , 1958 M. S c . ( F o r e s t r y ) U n i v e r s i t y o f B r i t i s h C o l u m b i a , 1967 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n t h e D e p a r t m e n t o f F o r e s t r y 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 t h e r e q u i r e d s t a n d a r d THE UNIVERSITY OF BRITISH A p r i l , 1971 COLUMBIA In present ing th i s thes i s in pa r t i a l f u l f i l m e n t of the requirements for an advanced degree at the Un ivers i ty of B r i t i s h Columbia, I agree that the L ibrary sha l l make i t f ree l y ava i l ab le for reference and study. I fu r ther agree that permission for extens ive copying of th i s thes i s fo r s cho la r l y purposes may be granted by the Head of my Department or by h i s representat ives . It i s understood that copying or pub l i ca t i on o f th i s thes i s f o r f i nanc i a l gain sha l l not be allowed without my wr i t ten permiss ion. Department of The Un iver s i t y o f B r i t i s h Columbia Vancouver 8, Canada ABSTRACT The h y p o t h e s i s i s e x a m i n e d t h a t , among i n d i v i d u a l t r a c h e i d s o f c o n i f e r o u s woods, t h e w e i g h t s o f h o l o c e l l u l o s e and a l p h a - c e l l u l o s e s k e l e t o n s a r e a d i r e c t q u a n t i t a t i v e f u n c t i o n o f t h e i r l e n g t h , i n d e p e n d e n t o f s p e c i e s . A t o t a l o f 28 a n n u a l i n c r e m e n t s , r e p r e s e n t i n g wood f r o m n i n e c o n i f e r o u s g e n e r a and s e v e n f a m i l i e s , s e p a r a t e d i n t o e a r l y w o o d and l a t e w o o d f r a c t i o n s , was d e l i g n i f i e d w i t h p e r a c e t i c a c i d a n d s u b s e q u e n t r e d u c t i o n w i t h NaBH^. F u r t h e r r e d u c t i o n t o a l p h a - c e l l u l o s e f o l l o w e d f o r a p o r t i o n o f t h e h o l o c u l l u l o s e s k e l e t o n s . A b o u t 750 i n d i v i d u a l h o l o - a n d a l p h a - c e l l u l o s e t r a c - h e i d s k e l e t o n s w e r e m e a s u r e d f o r l e n g t h and w e i g h t . A s p e c i a l l y d e v e l o p e d q u a r t z u l t r a m i c r o - b a l a n c e , h a v i n g a w e i g h i n g r a n g e o f 0.06 t o 14 y g and a p r e c i s i o n ± 0.03 y g , was c o n s t r u c t e d and u s e d t o w e i g h ; i n d i v i d u a l t r a c h e i d s . S t a t i s t i c a l a n a l y s e s i n d i c a t e d a s i g n i f i c a n t p o s i t i v e c u r v i l i n e a r r e l a t i o n s h i p b e t w e e n l e g n t h o f t r a c h e i d and w e i g h t o f i t s c a r b o h y d r a t e f r a c t i o n . E s t i m a t e d v a r i a t i o n s a c c o u n t e d f o r i n h o l o c e l l u l o s e and a l p h a - c e l l u l o s e s k e l e t o n w e i g h t s , by t h e l e n g t h f a c t o r . ; a l o n e w e r e , r e s p e c t i v e l y , 91.9 and 95.7 p e r c e n t f o r p o o l e d d a t a o f a l l t h e s p e c i e s . No s i g n i f i c a n t d i f - f e r e n c e s i n h o l o c e l l u l o s e s k e l e t o n w e i g h t s w e r e e v i d e n t w i t h i n s p e c i e s f o r t h e same t r a c h e i d l e n g t h , w h e r e a s w e i g h t s o f a l p h a - c e l l u l o s e s k e l e t o n s w i t h i n s p e c i e s , a n d b o t h t h e h o l o - a n d a l p h a - c e l l u l o s e b e t w e e n s p e c i e s , d i f f e r e d s i g n i f i c a n t l y . R a d i a l v a r i a t i o n f o r s i n g l e t r a c h e i d w e i g h t s f o l l o w e d t r e n d s s i m i l a r t o t h o s e e s t a b l i s h e d by o t h e r s f o r s p e c i f i c g r a v i t y , and p e r c e n t a g e o f c e l l u l o s e b a s e d on g r o s s wood a n a l y - s e s . I n d i v i d u a l t r a c h e i d s o f j u v e n i l e wood had s i g n i f i c a n t l y l o w e r ( 1 % l e v e l ) a l p h a - c e l l u l o s e s k e l e t o n w e i g h t s t h a n t h o s e f r o m m a t u r e and o v e r m a t u r e wood, w h i l e d i f f e r e n c e s were non- s i g n i f i c a n t f o r h o l o c e l l u l o s e . O v e r m a t u r e wood t r a c h e i d s were s i g n i f i c a n t l y l i g h t e r ( c a r b o h y d r a t e s k e l e t o n w e i g h t ) t h a n t h o s e f r o m m a t u r e wood, f o r t h e same t r a c h e i d l e n g t h . D i f f e r e n c e s b etween e a r l y w o o d and l a t e w o o d were e x p l o r e d . F o r t h e same t r a c h e i d l e n g t h , b o t h e a r l y w o o d and l a t e w o o d t r a c - h e i d s c o n t a i n e d s i m i l a r amounts o f a l p h a - c e l l u l o s e , w h e r e a s t h e amount o f h o l o c e l l u l o s e p e r t r a c h e i d was h i g h e r i n l a t e w o o d . E x a m i n a t i o n o f c o m p r e s s i o n wood a l s o p r o v i d e d p o s i t i v e e v i d e n c e f o r t h e l e n g t h - w e i g h t r e l a t i o n s h i p i n t r a c h e i d s . When w e i g h t s o f c o m p r e s s i o n wood t r a c h e i d s were compared w i t h t h o s e o f r e g u l a r (normal) wood, no s i g n i f i c a n t d i f f e r e n c e s were a p p a r e n t f o r h o l o c e l l u l o s e t r a c h e i d s k e l e t o n s , whereas s i g n i f i - c a n t d i f f e r e n c e s were f o u n d f o r a l p h a - c e l l u l o s e . I t was c o n - c l u d e d t h a t , f o r t h e same t r a c h e i d l e n g t h , c o m p r e s s i o n wood t r a c h e i d s may.have a l o w e r amount o f a l p h a - c e l l u l o s e t h a n t h o s e f r o m m a t u r e (normal) wood, b u t a h i g h e r amount t h a n t h o s e f r o m j u v e n i l e wood. Changes i n D o u g l a s - f i r t r a c h e i d w e i g h t s were s t u d i e d i n wood f o r m e d b e f o r e and f o l l o w i n g t r e e f e r t i l i z a t i o n . V a r i a t i o n s , f o r t h e most p a r t , were f o u n d t o be a s s o c i a t e d w i t h changes i n t r a c h e i d l e n g t h . Q u a l i t a t i v e d i f f e r e n c e s a t t r i - b u t a b l e t o t r e a t m e n t c o m p o s i t i o n ( U r e a v s . NPK v s . (NH^^SO^) were a l s o n o t e d , i n t h a t some t r e a t m e n t s r e s u l t e d i n l e s s w e i g h t o f a l p h a - c e l l u l o s e p e r u n i t l e n g t h o f t r a c h e i d , when compared w i t h n o r m a l wood t r a c h e i d s . T h i s r e d u c t i o n i n c e l l u l o s e f r a c - t i o n was s u g g e s t e d as a p o s s i b l e f a c t o r f o r d i f f e r e n c e s o b s e r v e d i n g r o s s wood s p e c i f i c g r a v i t y i n wood o f some f e r t i l i z e d t r e e s . R e s u l t s were c o m p a t i b l e w i t h t h e p r o p o s e d h y p o t h e s i s . i v TABLE OF CONTENTS Page ABSTRACT i TABLE OF CONTENTS i v L I S T OF TABLES vi LIST,OF FIGURES v i i i L I S T OF APPENDICES x ACKNOWLEDGEMENT x i CHAPTER I . INTRODUCTION 1 I I . LITERATURE REVIEW 4 T r a c h e i d M o r p h o l o g y 4 T r a c h e i d L e n g t h V a r i a t i o n 11 D i m e n s i o n a l R e l a t i o n s h i p s i n T r a c h e i d s 15 C h e m i c a l C o m p o s i t i o n and R e c o v e r y o f C a r b o h y d r a t e F r a c t i o n 19 E s t i m a t e s o f C a r b o h y d r a t e F r a c t i o n and W e i g h t i n R e l a t i o n t o C e l l u l a r E l e m e n t s 26 E f f e c t s o f F e r t i l i z a t i o n on Wood P r o p e r t i e s and C h e m i c a l C o m p o s i t i o n 29 I I I . MATERIALS AND METHODS 3 5 S e l e c t i o n o f M a t e r i a l 35 Wood Sample and T r a c h e i d S k e l e t o n P r e p a r a t i o n 37 A n a l y s i s o f T r a c h e i d S k e l e t o n s 40 1. S a m p l i n g c r i t e r i o n 40 2. T r a c h e i d l e n g t h 41 3. T r a c h e i d w e i g h t 42 V CHAPTER Page S t a t i s t i c a l A n a l y s e s 45 I V . RESULTS AND DISCUSSION 47 R e l a t i o n s h i p B e t w e e n T r a c h e i d L e n g t h a nd W e i g h t 48 1. V a r i a t i o n w i t h i n s p e c i e s 48 2. V a r i a t i o n b e t w e e n s p e c i e s 52 3. V a r i a t i o n w i t h i n c r e a s i n g age 55 4. V a r i a t i o n b e t w e e n r e g u l a r ( n o r m a l ) and c o m p r e s s i o n wood 60 5. . V a r i a t i o n b e t w e e n e a r l y w o o d a n d - l a t e w o o d 64 E v i d e n c e f o r a Common R e l a t i o n s h i p i n C o n i f e r s 68 A p p l i c a b i l i t y o f t h e R e l a t i o n s h i p 72 Some P r a c t i c a l C o n s i d e r a t i o n s 82 V. RECOMMENDATIONS FOR FURTHER RESEARCH 8 6 V I . CONCLUSION 8 8 LITERATURE CITED 91 TABLES 105 FOREWORD TO FIGURES 120 FIGURES 121 APPENDICES 139 v i L I S T OF TABLES NUMBER P a g e I . S i n g l e t r a c h e i d ( h o l o c e l l u l o s e s k e l e t o n ) w e i g h t s and l e n g t h s f o r n i n e c o n i f e r o u s woods 105 I I . S i n g l e t r a c h e i d ( a l p h a - c e l l u l o s e s k e l e t o n ) w e i g h t s and l e n g t h s f o r n i n e c o n i f e r o u s woods 107 I I I . S i n g l e t r a c h e i d ( h o l o c e l l u l o s e s k e l e t o n ) w e i g h t s a n d - l e n g t h s f o r two D o u g l a s - f i r t r e e s 109 I V . S i n g l e t r a c h e i d ( a l p h a - c e l l u l o s e s k e l e t o n ) w e i g h t s and l e n g t h s f o r two D o u g l a s - f i r t r e e s 110 V. A n a l y s i s m o d e l (A) and (B) r e s u l t s o f m u l t i p l e c u r v i l i n e a r c o v a r i a n c e a n a l y s e s ; d i f f e r e n c e s among s p e c i e s ( h o l o c e l l u l o s e s k e l e t o n s ) . ( A d j u s t e d f o r d i f f e r e n c e s i n t r a c h e i d l e n g t h ) 111 V I . A n a l y s i s m o d e l (A) and (B) r e s u l t s o f m u l t i p l e c u r v i l i n e a r c o v a r i a n c e a n a l y s e s ; d i f f e r e n c e s among s p e c i e s ( a l p h a - c e l l u l o s e s k e l e t o n s ) . ( A d j u s t e d f o r d i f f e r e n c e s i n t r a c h e i d l e n g t h ) 112 V I I . M u l t i p l e c u r v i l i n e a r c o v a r i a n c e a n a l y s i s f o r t r a c h e i d w e i g h t s ( h o l o c e l l u l o s e s k e l e t o n s ) i n j u v e n i l e , m a t u r e and o v e r m a t u r e wood o f a 5 0 0 ^ y e a r - o l d D o u g l a s - f i r t r e e . . ( A d j u s t e d f o r d i f f e r e n c e s i n t r a c h e i d l e n g t h ) 113 V I I I . M u l t i p l e c u r v i l i n e a r c o v a r i a n c e a n a l y s i s f o r t r a c h e i d w e i g h t s ( a l p h a - c e l l u l o s e s k e l e t o n s ) i n j u v e n i l e , m a t u r e a n d o v e r m a t u r e wood o f a 5 0 0 - y e a r - o l d D o u g l a s - f i r t r e e . ( A d j u s t e d f o r d i f f e r e n c e s i n t r a c h e i d l e n g t h ) 114 I X . A n a l y s i s o f c o v a r i a n c e a n d a d j u s t e d mean v a l u e s o f e a r l y w o o d and l a t e w o o d t r a c h e i d w e i g h t s i n s i x D o u g l a s - f i r t r e e s . ( A d j u s t e d f o r d i f - f e r e n c e s i n t r a c h e i d l e n g t h ) 115 X. A n a l y s i s o f • c o v a r i a n c e a n d a d j u s t e d mean v a l u e s o f e a r l y w o o d a n d l a t e w o o d t r a c h e i d w e i g h t s i n n i n e c o n i f e r o u s s p e c i e s . ( A d j u s t e d f o r d i f - f e r e n c e s i n t r a c h e i d l e n g t h ) 116 X I . S e l e c t e d wood a n d t r a c h e i d p r o p e r t i e s o f t h r e e f e r t i l i z e d D o u g l a s - f i r t r e e s 117 v i i NUMBER X I I . Page R e s u l t s o f m u l t i p l e c u r v i l i n e a r c o v a r i a n c e a n a l y s e s ; d i f f e r e n c e s w i t h i n s p e c i e s i n t r e a t e d and u n t r e a t e d D o u g l a s - f i r . ( A d j u s t e d f o r d i f f e r e n c e s i n t r a c h e i d l e n g t h ) 118 v i i i L I S T OF FIGURES NUMBER P a g e 1. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s , f r o m A r a u c a r i a c u n n i n g h a m i i 121 2. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) a nd l e n g t h f o r s a m p l e s ~ f r o m S e q u o i a s e m p e r v i r e n s 122 3. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) a nd l e n g t h f o r samples^" f r o m P i n u s l a m b e r t i a n a . 123 4. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) a nd l e n g t h f o r samples"'.from P o d o c a r p u s d a c r y d i o i d e s 124 5. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) a nd l e n g t h f o r samples.: f r o m P i c e a s i t c h e n s i s 125 6. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B)" s k e l e t o n s ) and l e n g t h f o r s a m p l e s f f r o m T a x u s b r e v i f o l i a , 126 7. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r samples-" f r o m J u n i p e r u s v i r g i n i a n a 127 8. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) a nd l e n g t h f o r samples." f r o m C e p h a l o t a x u s w i l s o n i a n a 128 9. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) a n d a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s f r o m P s e u d o t s u g a m e n z i e s i i 129 i x NUMBER P a g e 10. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e . (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s a n d - l e n g t h f o r s a m p l e s f r o m n i n e c o n i f e r o u s s p e c i e s . ( L e a s t - s q u a r e s f i t t e d l i n e ) 130 1 1 . R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s and l e n g t h f o r s a m p l e s f r o m n i n e c o n i f e r o u s s p e c i e s . (The l i n e o f b e s t f i t c o n d i t i o n e d t o p a s s t h r o u g h t h e o r i g i n ) 131 12. T r a c h e i d w e i g h t , mean t r a c h e i d l e n g t h a n d s p e c i f i c g r a v i t y p a t t e r n s , a c r o s s t h e b u t t l o g o f a 500- y e a r - o l d D o u g l a s - f i r t r e e . ( E a c h p o i n t r e p r e - s e n t s t h e a v e r a g e t r a c h e i d w e i g h t o r l e n g t h f o r t h e i n c r e m e n t . S p e c i f i c g r a v i t y - v a l u e s f r o m Kennedy a nd W a r r e n ( 7 0 ) ) 132 13. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r c o m p r e s s i o n wood and n o r m a l wood o f D o u g l a s - f i r . 133 14. T r a c h e i d w e i g h t , i n c r e m e n t w i d t h a n d mean t r a c h e i d l e n g t h p a t t e r n s a c r o s s s i x c o n s e c u t i v e i n c r e - m e nts o f a f e r t i l i z e d (NH4NO3) D o u g l a s - f i r w h i c h p r o d u c e d c o m p r e s s i o n wood 4 y e a r s a f t e r f i r s t f e r t i l i z a t i o n . ( E a c h p o i n t r e p r e s e n t s t h e a v e r a g e t r a c h e i d w e i g h t o r l e n g t h f o r t h e i n c r e m e n t ) 134 15. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) a n d a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h i n U r e a t r e a t e d a n d n o r m a l D o u g l a s - f i r 135 16. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) a n d l e n g t h i n NPK t r e a t e d a n d n o r m a l D o u g l a s - f i r 136 17. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e (A) a n d a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h i n ( N B ^ ^ S O ^ t r e a t e d and n o r m a l D o u g l a s - f i r 137 18. R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o - c e l l u l o s e ,-(A) a n d a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h i n NH4NO3 t r e a t e d a n d n o r m a l D o u g l a s - f i r 13 8 X L I S T OF APPENDICES NUMBER Page I . D e s c r i p t i o n o f g r o w t h i n c r e m e n t s i n c l u d e d i n t h e s t u d y 13 9 I I . Summary o f h o l o c e l l u l o s e and a l p h a - c e l l u l o s e y i e l d s and e s t i m a t e s o f l i g n i n i n some o f t h e s a m p l e s s t u d i e d ( b a s e d o n o v e n d r y , e x t r a c t i v e - f r e e wood) 141 I I I . M o i s t u r e c o n t e n t o f t h e h o l o c e l l u l o s e a n d a l p h a - c e l l u l o s e p u l p s a f t e r c o n d i t i o n i n g i n t h e CTH Room f o r o v e r one month 142 x i ACKNOWLEDGEMENT The w r i t e r a c k n o w l e d g e s t h e c o n t i n u o u s g u i d a n c e , i n s p i r a t i o n and c o n s t r u c t i v e c r i t i c i s m g i v e n t o h i m d u r i n g h i s e n t i r e s t a y a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , by h i s s u p e r v i s o r , D r . R. W. W e l l w o o d . W i t h o u t t h e i n t e r e s t shown by h i m t h e t h e s i s m i g h t n o t h a v e b e e n u n d e r t a k e n o r c o m p l e t e d . The h e l p a n d k i n d n e s s o f D r . J.W. W i l s o n i s acknow- l e d g e d , n o t o n l y f o r h i s many u s e f u l s u g g e s t i o n s d u r i n g t h e p l a n n i n g a n d e x p e r i m e n t a l p h a s e s o f t h e t h e s i s , b u t a l s o f o r h i s t r u l y e n l i g h t e n i n g i n s p i r a t i o n , e n c o u r a g e m e n t and p e r s o n a l c o n c e r n . D r . A. K o z a k h e l p e d t h e w r i t e r i n c l a r i f y i n g many q u e s t i o n s c o n c e r n i n g s t a t i s t i c a l a n a l y s e s . H i s a d v i c e i s g r a t e f u l l y a c k n o w l e d g e d . The t h e s i s was r e v i e w e d by members o f t h e a d v i s o r y c o m m i t t e e c o n s i s t i n g o f D r s . R.W. .Kennedy.,. A. .-Kozak, RvW.-. W e l l w o o d , J.W. W i l s o n , . J . W o r r a l l and D.J. W o r t . F o r t h e i r v a l u a b l e s u g g e s t i o n s , c o m m e n t s a n d c o n s t r u c t i v e c r i t i c i s m t h e w r i t e r i s i n d e b t e d . Many h e l p f u l s u g g e s t i o n s f r o m D r s . O.H. L o w r y , W a s h i n g t o n U n i v e r s i t y S c h o o l o f M e d i c i n e , S t . L o u i s , D.S. S k e n e , C a b o t F o u n d a t i o n f o r B o t a n i c a l R e s e a r c h , H a r v a r d F o r e s t , P e t e r - sham, B.S. Wenger, D e p a r t m e n t o f Anatomy, S c h o o l o f M e d i c i n e , U n i v e r s i t y o f S a s k a t c h e w a n , and t h e t i m e and t a l e n t s o f many x i i persons on the F a c u l t y of F o r e s t r y t e c h n i c a l s t a f f , Messrs. U. Rumma, K. Apt, and G. Bohnenkamp, made p o s s i b l e develop- ment of the q u a r t z u l t r a - m i c r o b a l a n c e , an i n t e g r a l p a r t of the s tudy. D i s c u s s i o n s with the f o l l o w i n g c o n t r i b u t e d g r e a t l y , i n understanding some of the problems i n v o l v e d h e r e i n and t h e i r advice i s a p p r e c i a t e d . Dr. R.W. Kennedy Department of F i s h e r i e s and F o r e s t r y , Canadian F o r e s t r y S e r v i c e F o r e s t Products Laboratory Vancouver. Dr. L. Paszner F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia. Dr. J . W o r r a l l F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia. Dr. D.J. Wort Department of Botany, U n i v e r s i t y of B r i t i s h Columbia.- The w r i t e r s i n c e r e l y thanks Miss L. Cowdell and Mrs. K. Hejjas f o r w r i t i n g the computer programs; Mrs. M. Lambden f o r d r a f t i n g work; Messrs H . J . Cho and C.P. Chen, Graduate Students, f o r t e c h n i c a l a s s i s t a n c e i n the p r e p a r a t i o n of pulp samples; the N a t i o n a l Research C o u n c i l of Canada and the U n i v e r s i t y . o f B r i t i s h Columbia f o r f i n a n c i a l a s s i s t a n c e . F i n a l l y , he must express h i s s p e c i a l a p p r e c i a t i o n to h i s w i f e , Ratna, f o r her encouragement and monumental pat i e n c e . CHAPTER I INTRODUCTION Much c u r r e n t r e s e a r c h i n wood and p u l p s c i e n c e e m p h a s i z e s v a r i a b i l i t y a t t h e c e l l u l a r l e v e l . The p r i n c i p a l c e l l u l a r e l e m e n t s o f c o n i f e r o u s woods a r e l o n g i t u d i n a l t r a c h e i d s w h i c h make up as much as 90 p e r c e n t by volume o r 98 p e r c e n t by w e i g h t o f t h e o v e n d r y m a t e r i a l . S i n c e t h e t r a c h e i d i s t h e b a s i c u n i t o f s u c h woods, i t s i s o l a t i o n and s t u d y s h o u l d a s s i s t i n u n d e r s t a n d i n g and i n t e r p r e t i n g t h e b i o l o g i c a l a c t i v i t y and p r o d u c t b e h a v i o u r o f wood. V a r i a b i l i t y i n wood w i t h i n and between s p e c i e s has been s t u d i e d e x t e n s i v e l y a t t h e g r o s s l e v e l . S i m i l a r l y , v a r i a - t i o n w i t h i n a s i n g l e stem has b e e n s t u d i e d w i t h r e s p e c t t o r a d i a l and h e i g h t p a t t e r n s , as w e l l as wood zones (corewood, h e a r t w o o d and sapwood). L e s s f r e q u e n t l y , s e p a r a t e e a r l y w o o d and l a t e w o o d segments have been a n a l y z e d f o r d i f f e r e n c e s i n p h y s i c a l b e h a v i o u r . A t s t i l l a n o t h e r l e v e l o f i n v e s t i g a t i o n , t h e v a r i a b i l i t y i n p h y s i c a l and c h e m i c a l p r o p e r t i e s w i t h i n i n d i v i d u a l g r o w t h i n c r e m e n t s has b e e n examined by i n t r a - i n c r e - m e n t a l a n a l y s e s . F u r t h e r i n t e n s i v e s t u d y o c c u r r e d a t t h e i n d i v i d u a l c e l l l e v e l , i n p a r t i c u l a r c e l l w a l l o r g a n i z a t i o n and c h e m i c a l c o m p o s i t i o n i n c l u d i n g s p a t i a l d i s t r i b u t i o n o f t h e c o n s t i t u e n t s w i t h i n c e l l w a l l s . 2. E v e n t h o u g h c h e m i c a l c o n s t i t u e n t s and p h y s i c a l p r o p e r - t i e s o f wood have b e e n i n t e n s i v e l y s t u d i e d , r e l a t i v e l y l i t t l e i s known a b o u t t h e amount o f c a r b o h y d r a t e f r a c t i o n p e r i n d i - v i d u a l t r a c h e i d , and p o s s i b l e r e l a t i o n t o an e a s i l y m e a s u r e d a n a t o m i c a l c h a r a c t e r i s t i c , f o r example, l e n g t h . Though s u c h i n f o r m a t i o n has b e e n d e d u c e d by i n d i r e c t methods s u c h as f i b r e c o a r s e n e s s , o r t i s s u e f r a c t i o n s , t h e s e methods have an i n h e r e n t weakness i n t h a t t h e e s t i m a t e o f i n d i v i d u a l c e l l u l a r v a r i a t i o n i s n o t p o s s i b l e . S c i e n t i s t s i n v a r i o u s segments o f t h e wood i n d u s t r i e s a r e e m p h a s i z i n g methods o f d e f i n i n g wood q u a l i t y . T h e s e e f f o r t s h a v e l e a d t o a s e a r c h f o r an a n a t o m i c a l b a s i s f o r e x p r e s s i o n o f wood q u a l i t y . D i m e n s i o n s o f t r a c h e i d s and t h e i r c a r b o h y d r a t e c o n t e n t a r e i m p o r t a n t b e c a u s e e a c h i s g e n e r a l l y known t o be c l o s e l y r e l a t e d t o g r o s s wood c h a r a c t e r i s t i c s t h a t a r e u s e d as q u a l i t y i n d i c a t o r s , s u c h as d e n s i t y , and v a r i o u s s t r e n g t h p r o p e r t i e s . A l s o , t h e i n f l u e n c e o f t r a c h e i d l e n g t h and s t r e n g t h o f i n d i v i d u a l f i b r e 1 on p u l p and p a p e r p r o p e r t i e s h as been e s t a b l i s h e d . The s i g n i f i c a n c e o f c e l l u l o s e c o n t e n t on t e n s i l e s t r e n g t h o f wood has b e e n c o n f i r m e d by v a r i o u s w o r k e r s . C e r t a i n l y , a t h o r o u g h knowledge o f t h e r e l a t i o n s h i p between l e n g t h o f f i b r e and i t s c a r b o h y d r a t e amount, w i t h i n and between s p e c i e s , i s n e c e s s a r y i n o r d e r t o more f u l l y e v a l u a t e and u n d e r s t a n d g r o s s wood "*"The word ' f i b r e ' and ' t r a c h e i d ' a r e u s e d i n t e r - c h a n g e a b l y i n t h i s t h e s i s . p r o p e r t i e s and to more f u l l y u t i l i z e wood. In a d d i t i o n , know- ledge r e g a r d i n g the type of r e l a t i o n s h i p ( i . e . , l i n e a r or e x p o n e n t i a l ) , would be of importance to g e n e t i c i s t s who are endeavouring to manipulate wood c h a r a c t e r i s t i c s through t r e e b reeding work. Even more important, any gains i n t h i s funda- mental knowledge should be of g e n e r a l v a l u e to wood-based t e c h n o l o g i e s . The o b j e c t i v e s of t h i s t h e s i s are: 1. To t e s t the hypothesis t h a t , among i n d i v i d u a l t r a c h e i d s of c o n i f e r o u s woods, the weights of h o l o - c u l l u l o s e and a l p h a - c e l l u l o s e s k e l e t o n s are a d i r e c t q u a n t i t a t i v e f u n c t i o n of t h e i r l e n g t h , independent of s p e c i e s . 2. a. To apply the r e l a t i o n s h i p w i t h i n a s i n g l e s p e c i e s , D o u g l a s - f i r (Pseudotsuga m e n z i e s i i (Mirb.) Franco) by comparing r e g u l a r (normal) wood with j u v e n i l e wood, overmature wood, compression wood, and wood formed w i t h i n t r e e s t r e a t e d w i t h chemical f e r - t i l i z e r s . b. To examine the r e l a t i o n s h i p w i t h i n increments by comparing i n d i v i d u a l carbohydrate s k e l e t o n s from earlywood and latewood. CHAPTER I I LITERATURE REVIEW D u r i n g t h e p a s t 40 t o 50 y e a r s d a t a r e g a r d i n g t h e a n a t o m i c a l s t r u c t u r e , p h y s i c a l p r o p e r t i e s , c h e m i c a l c o m p o s i t i o n , and p h y s i c a l f u n c t i o n i n g o f wood h a v e a c c u m u l a t e d i n e v e r - i n c r e a s i n g v o l u m e . S e v e r a l books have been w r i t t e n on t h e s u b j e c t w i t h i n r e c e n t y e a r s (45, 62, 72, 101, 137) and t h e s e have been s u p p l e m e n t e d by r e v i e w p a p e r s (19, 20, 25, 31, 34, 70, 127, 136, 153, 1 6 8 ) . I t w o u l d seem s u p e r f l u o u s t o r e - c o v e r t h e same g r o u n d . An a t t e m p t w i l l be made t o c o v e r o n l y t h o s e p o i n t s o f l i t e r a t u r e w h i c h a r e r e l a t e d d i r e c t l y t o t h e t h e s i s t o p i c . T r a c h e i d M o r p h o l o g y As s t a t e d by Wardrop ( 1 4 5 ) , t h e t e r m m o r p h o l o g y was i n t r o d u c e d o r i g i n a l l y by G o e t h e e a r l y i n t h e l a s t c e n t u r y t o d e f i n e t h a t f i e l d o f s t u d y c o n c e r n e d w i t h p l a n t and a n i m a l f o r m , as d i s t i n c t f r o m t h e s t u d y o f f u n c t i o n . W i t h t i m e , t h e s c o p e o f t h e t e r m has b e e n w i d e n e d t o i n c l u d e t h e s t u d y o f b o t h s t r u c t u r e and f o r m , and more r e c e n t l y t o i n c l u d e t h e s t u d y o f s t r u c t u r e e v e n a t t h e m o l e c u l a r l e v e l . T h e r e f o r e , d i s c u s s i o n i s c o n c e r n e d n o t o n l y w i t h wood f i b r e e x t e r n a l g e o m e t r y b u t a l s o w i t h i n t e r n a l s t r u c t u r e (where a p p r o p r i a t e ) and, b r i e f l y w i t h t h e p h y s i o l o g y o f wood f o r m a t i o n . 5. The p r o d u c t i o n o f wood r e s u l t s f r o m a c t i v i t i e s o f t h e v a s c u l a r cambium w h i c h becomes c i r c u m f e r e n t i a l l y c o n t i n u o u s a s h o r t d i s t a n c e b e l o w t h e g r o w i n g t i p o f t h e m a i n s t e m and t h e b r a n c h e s o f a t r e e ( 4 5 ) . T h e r e a r e two c e l l t y p e s i n t h e v a s c u l a r cambium: 1) f u s i f o r m i n i t i a l s ; and 2) r a y i n i t i a l s . F u s i f o r m i n i t i a l s a r e e l o n g a t e d c e l l s w i t h t a p e r i n g e n d s , w h e r e a s t h e r a y i n i t i a l s a r e much s m a l l e r and a l m o s t i s o d i a m e t r i c i n s h a p e ( 1 9 ) . The f o r m e r i n i t i a l s g i v e r i s e t o a l l t h e v e r t i c a l e l o n g a t e e l e m e n t s o f t h e x y l e m and p h l o e m , w h e r e a s t h e r a y i n i t i a l s p r o d u c e a l l e l e m e n t s o f t h e t r a n s v e r s e o r r a y s y s t e m o f t h e x y l e m and p h l o e m ( 4 5 ) . Two t y p e s o f c e l l d i v i s i o n o c c u r i n t h e v a s c u l a r cambium o f c o n i f e r s ; p e r i c l i n a l o r a d d i t i v e a nd a n t i c l i n a l o r m u l t i p l i c a t i v e . P e r i c l i n a l d i v i s i o n s a d d t o t h e x y l e m and p h l o e m h e n c e b r i n g i r . a b o u t r a d i a l g r o w t h , w h i l e t h e l a t t e r t y p e r e s u l t i n m u l t i p l i c a t i o n o f f u s i f o r m c a m b i a l c e l l s w i t h c o n s e - q u e n t a c c o m m o d a t i o n t o c i r c u m f e r e n t i a l e x p a n s i o n ( 8 ) . B e c a u s e t h e f u s i f o r m i n i t i a l s g i v e r i s e t o a l l t h e v e r t i c a l e l o n g a t e e l e - m ents o f t h e x y l e m and p h l o e m , t h e i r i n i t i a l a n d f i n a l d i m e n s i o n s , and t h e many p h y s i o l o g i c a l f a c t o r s a f f e c t i n g t h e d i m e n s i o n s , a r e o f g r e a t i m p o r t a n c e t o t h e k i n d a n d q u a l i t y o f wood p r o d u c e d . T h i s i s p a r t i c u l a r l y i m p o r t a n t t o c o n i f e r s s i n c e t h e l e n g t h o f d e r i v e d v a s c u l a r e l e m e n t s i s t r a c e a b l e b a c k t o c a m b i a l i n i t i a l s 6 . f r o m w h i c h t r a c h e i d s were d e r i v e d , i . e . , p o s t - d i v i s i o n a l e l o n - g a t i o n o f d a u g h t e r c e l l s i s r e l a t i v e l y s l i g h t i n c o n i f e r s (2, .8) . The p a t t e r n o f s e c o n d a r y w a l l f o r m a t i o n i n f i b r e s has b e e n e x t e n s i v e l y s t u d i e d by A u s t r a l i a n w o r k e r s , and W ardrop, i n two r e c e n t a r t i c l e s , has summarized t h e work i n t h i s a r e a (143, 1 4 4 ) . The d i s c u s s i o n t h a t f o l l o w s i s a b r i e f s y n t h e s i s o f t h e e x c e l l e n t and e x t e n s i v e work o f Wardrop and h i s a s s o c i a t e s (143, 144, 145, 146, 147, 1 4 8 ) . D e v e l o p m e n t o f t r a c h e i d s f r o m t h e cambium o f c o n i f e r stems i n v o l v e s f o u r p h a s e s : 1) c e l l d i v i s i o n i n t h e cambium; 2) e n l a r g e m e n t o f t h e d a u g h t e r c e l l s t o t h e d imen- s i o n s o f t h e m a t u r e c e l l ; 3) t h e d e v e l o p m e n t o f t h e . s e c o n d a r y w a l l ; and 4) l i g n i f i c a t i o n ( 1 4 7 ) . D a u g h t e r c e l l s , f o r m e d as a r e s u l t o f c e l l d i v i s i o n i n t h e cam- bium, i m m e d i a t e l y u n d e r g o c o n s i d e r a b l e d i m e n s i o n a l change, i n c r e a s i n g r a d i a l l y by 200 t o 500 p e r c e n t and i n l e n g t h by 10 t o 20 p e r c e n t o r more. D e v e l o p m e n t o f s e c o n d a r y w a l l b e g i n s , i n p a r t , b e f o r e d i m e n s i o n a l c h a n g e s o f d i f f e r e n t i a t i o n a r e c o m p l e t e , f i r s t n e a r t h e m i d d l e o f t h e d i f f e r e n t i a t i n g t r a c h e i d s t h e n e x t e n d s t o w a r d t h e i r t i p s . D u r i n g t h e p r o g r e s - s i v e d e v e l o p m e n t o f t h e s e c o n d a r y w a l l , c o n t a c t between t h e 7. p r i m a r y w a l l and t h e p r o t o p l a s t i s m a i n t a i n e d by a m i n u t e t e r m i n a l c a n a l b etween t h e lumen and t h e p r i m a r y w a l l ( 1 4 7 ) . Wardrop and D a d s w e l l (146) r e p o r t e d t h a t t h e t h i c k e n i n g o f t h e c e l l w a l l i n m a t u r e t r a c h e i d s was o f t e n l e s s a t t h e t i p s t h a n i n t h e m i d d l e . T h i s o b s e r v a t i o n , b e s i d e s s u p p o r t i n g t h e p r o - g r e s s i v e d e v e l o p m e n t o f s e c o n d a r y w a l l f r o m t h e m i d d l e t o w a r d s t h e c e l l t i p , a l s o i n d i c a t e s t h e p o s s i b i l i t y t h a t t h e t i p s p e r s e s h o u l d w e i g h : l e s s compared t o a s i m i l a r p o r t i o n o f u n i t l e n g t h f r o m t h e t r a c h e i d b ody. The v a r i o u s f a c t o r s w h i c h i n f l u e n c e t h e e x t e n t o f c e l l e n l a r g e m e n t and w a l l d e v e l o p m e n t f o l l o w i n g c e l l d i v i s i o n a r e s t i l l n o t f u l l y u n d e r s t o o d . C e l l e n l a r g e m e n t a p p a r e n t l y i s a f u n c t i o n o f c e l l v a c u o l a t i o n and i t has b e e n s u g g e s t e d t h a t t h i s i s r e l a t e d t o t h e w a t e r r e g i m e o f t h e t r e e s a t t h e t i m e t h a t c e l l e n l a r g e m e n t o c c u r s ( 1 4 ) . As n o t e d by Wardrop (148) : I n t h e l i v i n g c o n d i t i o n t h e c e l l s o f t h e cambium e x i s t i n a s t a t e o f t u r g o r and t h e c a p a c i t y o f t h e c e l l s t o t a k e up w a t e r i s de- t e r m i n e d by t h e d i f f e r e n c e b etween t h e a c t u a l p r e s s u r e m u t u a l l y e x e r t e d on one a n o t h e r by t h e w a l l and p r o t o p l a s t and t h e f u l l ( p o t e n - t i a l ) p r e s s u r e a v a i l a b l e t h r o u g h o s m o t i c a c t i v i t y o f s u b s t a n c e s i n t h e c e l l s a p . C e l l e l o n g a t i o n o f f u s i f o r m i n i t i a l s i s a c h i e v e d by i n t r u s i v e g r o w t h , i . e . , t i p s o f t h e e l o n g a t i n g c e l l e x t e n d a t one o r b o t h e n d s , between t h e o t h e r d e v e l o p i n g c e l l s (2, 9, 45, 1 0 1 ) . A c c o r d i n g t o c u r r e n t c o n c e p t s , r a d i a l e x p a n s i o n o f t h e t r a c h e i d i s r e g u l a t e d by a u x i n e m a n a t i n g f r o m g r o w t h c e n t e r s 8. i n t h e crown (77, 8 0 ) . I n a d d i t i o n , R o e l o f s e n (115) f o r m u l a t e d t h e m u l t i n e t t h e o r y f o r t h e g r o w t h o f t h e p r i m a r y w a l l . T h i s t h e o r y i m p l i e s t h a t t h e i n n e r m o s t p a r t o f t h e w a l l c o n s i s t s o f b u n d l e s o f more o r l e s s t r a n s v e r s e l y o r i e n t e d m i c r o f i b r i l s . D u r i n g c e l l e x p a n s i o n , t h e f i b r i l s a r e p a s s i v e l y d i s p l a c e d and s l i p a l o n g e a c h o t h e r as t h i s membrane s h e a t h i s moved o u t w a r d by t h e a p p o s i t i o n o f new w a l l m a t e r i a l , and t h e s t r a n d s become s u b s e q u e n t l y o r i e n t e d i n t h e d i r e c t i o n o f g r o w t h . However, t h e m u l t i n e t t h e o r y does n o t o f f e r an e x p l a n a t i o n f o r t h e t r a n s v e r s e o r i e n t a t i o n o f t h e f i b r i l s p r i o r t o c e l l w a l l e x p a n s i o n . From t h e s t a n d p o i n t o f i t s d e v e l o p m e n t , t h e m a t u r e c o n i f e r t r a c h e i d c o n s i s t s o f two s t r u c t u r e s : t h e p r i m a r y w a l l and t h e s e c o n d a r y w a l l . The p r e s e n t l y a c c e p t e d c o n c e p t and t e r m i n o l o g y o f c e l l w a l l s t r u c t u r e . w a s d e v e l o p e d by K e r r and B a i l e y ( 6 7 ) . They d i v i d e d t h e c e l l w a l l i n t o t h e p r i m a r y w a l l ( c a m b i a l w a l l ) and t h e s e c o n d a r y w a l l ( s e c o n d a r y t h i c k e n i n g l a i d down a f t e r t h e c e l l has r e a c h e d i t s f i n a l s i z e and s h a p e ) . The s e c o n d a r y w a l l c o n s i s t s o f t h r e e l a y e r s — a n o u t e r l a y e r , a m i d d l e S^r and an i n n e r S^--of d i f f e r i n g o p t i c a l p r o p e r t i e s ( 1 4 5 ) . T h i s o p t i c a l h e t e r o g e n e i t y has b e e n a t t r i b u t e d t o d i f - . f e r e n c e s i n h e l i c a l o r i e n t a t i o n o f t h e c e l l u l o s e m i c e l l a r s t r a n d s i n t h e t h r e e l a y e r s (143, 1 4 4 > r 1 4 5 ) . I n c o n i f e r s , t h e i n n e r m o s t l a y e r may be s c u l p t u r e d i n t h e f o r m o f s m a l l e x c r e s c e n c e s (wart s t r u c t u r e s ) (85, 1 4 5 ) , p o s s i b l y f o r m e d f r o m p r o t o p l a s m i c d e b r i s . T h e s e a s p e c t s h a v e been e x t e n s i v e l y 9. r e v i e w e d by Wardrop and h i s a s s o c i a t e s who d e v e l o p e d a s c h e m a t i c model o f t h e c e l l w a l l (142, 143, 1 4 4 ) . I n a d d i t i o n , D u n n i n g (36) r e c e n t l y d e v e l o p e d a.new method f o r m i c r o d i s s e c t i o n and e l e c t r o n m i c r o s c o p y o f s i n g l e wood p u l p f i b r e s , and p r o v i d e d a d d i t i o n a l i n f o r m a t i o n on c e l l w a l l o r g a n i z a t i o n , i n p a r t i c u l a r t h e l a y e r . From s t u d i e s o f Jayme and F e n g e l (63) and s e v e r a l o t h e r w o r k e r s r e f e r r e d t o by them, i t i s u n d e r s t o o d t h a t , f o r b o t h e a r l y w o o d and l a t e w o o d , t h e S2 l a y e r . i s b y . f a r t h e t h i c k e s t and p r o b a b l y a c c o u n t s f o r 80 t o 95 p e r c e n t o f t h e volume o r w e i g h t o f t h e f i b r e . V a r i a t i o n s do o c c u r i n t h e w a l l o r g a n i z a t i o n , as i n t h e c a s e o f r e a c t i o n wood anatomy, and i t i s r e p o r t e d t h a t e v e n i n n o r m a l wood some P i c e a spp. l a c k an l a y e r (53) . Compres- s i o n wood t r a c h e i d s d i f f e r i n c r o s s s e c t i o n f r o m n o r m a l ones i n t h e i r t y p i c a l r o u n d e d a p p e a r a n c e and i n t e r c e l l u l a r s p a c e s . They u s u a l l y h a v e ,a w i d e r l a y e r and numerous r e g u l a r r a d i a l d i s c o n t i n u i t i e s . i n t h e S^r as w e l l as a much g r e a t e r m i c r o - f i b r i l l a r a n g l e t h a n n o r m a l wood t r a c h e i d s . I n a d d i t i o n , com- p r e s s i o n wood t r a c h e i d s c o n t a i n an e x t r a l a y e r o f l i g n i n l o c a t e d between and ( 1 3 6 ) . A c c o r d i n g t o T i m e l l (13 6 ) , t e n s i o n wood o f t e n l a c k s one o r s e v e r a l o f t h e t h r e e s e c o n d a r y w a l l l a y e r s and f r e q u e n t l y p o s s e s s e s an a d d i t i o n a l w a l l l a y e r , d e p o s i t e d n e x t t o t h e lumen, t h e s o - c a l l e d g e l a t i n o u s l a y e r . T h i s c o n t a i n s l i t t l e i f any l i g n i n and c o n s i s t s l a r g e l y o f h i g h l y c r y s t a l l i n e m i c r o f i b r i l s o f c e l l u l o s e , o r i e n t e d p a r a l l e l t o t h e f i b r e a x i s "(136). S i m i l a r l y , t h e m o r p h o l o g y o f t r a c h e i d s , 10. i . e . , d i m e n s i o n s and t h i c k n e s s o f w a l l l a y e r s , d i f f e r s b e t w e e n e a r l y w o o d and l a t e w o o d o f t h e same g r o w t h i n c r e m e n t ( 1 3 7 ) . T h e r e i s a n i n d i c a t i o n , h o w e v e r , t h a t t h e c r o s s - s e c t i o n a l c e l l w a l l a r e a o f e a r l y w o o d a nd o f l a t e w o o d t r a c h e i d s may r e m a i n c o n s t a n t i n c o n i f e r s (14, 6 8 ) . A c c o r d i n g t o W a r d r o p (143) and as s u m m a r i z e d by S t e w a r t ( 1 2 7 ) , t h e c e l l w a l l c h e m i c a l c o n s t i t u e n t s may be c l a s s i f i e d i n t o t h r e e g r o u p s : 1) t h e m a t r i x s u b s t a n c e s ; 2) . t h e f r a m e w o r k s u b s t a n c e s ; a nd 3) t h e e n c r u s t i n g s u b s t a n c e s . The f r a m e w o r k ( c e l l u l o s i c ) a n d m a t r i x ( n o n - c e l l u l o s i c ) p o l y s a c c h a r i d e s a r e d e p o s i t e d b e f o r e t h e p r i m a r y ( l i g n i n ) a n d s e c o n d a r y ( e x t r a c t i v e s ) e n c r u s t i n g s u b s t a n c e s . The d i s t r i b u t i o n o f t h e c h e m i c a l c o n s t i t u e n t s i s u n e v e n t h r o u g h o u t t h e c e l l w a l l . B a i l e y (1) r e p o r t e d t h e i s o - l a t i o n o f t h e m i d d l e l a m e l l a o f D o u g l a s - f i r by m i c r o d i s s e c t i o n and i n d i c a t e d t h a t i t c o n t a i n e d 71 p e r c e n t l i g n i n , a n d 14 p e r c e n t p e n t o s a n s . L a n g e and h i s ass o c i a t e . ' . (75) a p p l i e d a m i c r o - s p e c t r o p h o t o m e t r i c m e t h o d o n c r o s s s e c t i o n s o f wood s a m p l e s o f S w e d i s h s p r u c e ( P i c e a a b i e s (L.) K a r s t ) , and c o n f i r m e d t h e e a r l i e r f i n d i n g s o f B a i l e y . I n a d d i t i o n , i t was n o t e d t h a t t h e l i g n i n c o n c e n t r a t i o n e x t e r n a l t o t h e l a y e r i s 73 p e r c e n t , c o m p a r e d w i t h 16 p e r c e n t n e a r t h e l u m e n . B e r l y n and M a r k (11) r e v i e w e d t h i s a s p e c t a nd c o n c l u d e d t h a t a l t h o u g h m o s t o f t h e compound m i d d l e l a m e l l a , i s l i g n i r i ; - . t h i s d o e s n o t n e c e s s a r i l y - mean t h a t m o s t - o f t h e - l i g n i n ' (amount). :.is i n t h i s r e g i o n . 11. I n f o r m a t i o n on t h e s p a t i a l d i s t r i b u t i o n o f p o l y - s a c c h a r i d e s i n t h e c e l l w a l l i s l i m i t e d . The p r o p o r t i o n o f i n d i v i d u a l p o l y s a c c h a r i d e s i n e a c h c e l l w a l l l a y e r h a d b e e n d e d u c e d v e r y r o u g h l y by f o l l o w i n g m i c r o d i s s e c t i o n t e c h n i q u e s (90, 91) o r by m i c r o s p e c t r o p h o t o m e t r i c methods ( 7 4 ) . I n t h e i r r e c e n t book, P a n s h i n and-de Zeeuw (101) summarized t h e work i n t h i s a r e a as f o l l o w s : . . . l e s s t h a n 10 p e r c e n t o f t h e p r i m a r y w a l l i s c e l l u l o s i c ; i n t h e S ? l a y e r t h e c e l l u l o s e i n c r e a s e s t o more t h a n 50 p e r c e n t o f t h e m a t e r i a l i n t h a t c e l l w a l l l a y e r ; a r e d u c t i o n i n amount o f c e l l u l o s e - i s " e v i d e n t i n t h e S^. I t i s c l e a r f r o m t h e s e s t u d i e s t h a t c e l l u l o s e i s t h e m a j o r component i n t h e s e c o n d a r y w a l l , whereas l i g n i n and non- c e l l u l o s i c p o l y s a c c h a r i d e s a r e d o m i n a n t ' i n t h e p r i m a r y w a l l ( 1 4 5 ) . F u r t h e r m o r e , a c o m p a r i s o n o f t h e s e . v a l u e s w i t h t h o s e o f t h e l i g n i n i n d i c a t e t h a t p e r c e n t a g e o f l i g n i n i s r e - l a t e d i n v e r s e l y t o t h a t o f t h e t o t a l p o l y s a c c h a r i d e s ( 1 0 1 ) . Thus .the work done s o f a r has c e n t r e d m o s t l y o n t h e s p a t i a l d i s t r i b u t i o n . o f components i n t h e t r a c h e i d , i . e . , q u a l i t a t i v e a s p e c t s , b u t v e r y l i t t l e a t t e m p t has been made t o q u a n t i f y t h e v a r i o u s components p e r t r a c h e i d . T r a c h e i d L e n g t h V a r i a t i o n An e x t e n s i v e l i t e r a t u r e has a c c u m u l a t e d on t r a c h e i d l e n g t h v a r i a t i o n s b o t h w i t h i n and between t r e e s and on p r o b a b l e e f f e c t s o f t h e e x t e r n a l e n v i r o n m e n t . The p i o n e e r i n g work i n t h i s a r e a i s t r a c e a b l e t o S a n i o ' s f i r s t i n v e s t i g a t i o n s (4) o f t r a c h e i d l e n g t h v a r i a t i o n i n S c o t c h p i n e ( P i n u s s y I v e s t r i s L . ) . W i t h i n t h e p r e s e n t c e n t u r y t h e m a t t e r o f f i b r e l e n g t h has b e - come i m p o r t a n t t o t h e p u l p and p a p e r i n d u s t r y , w i t h a c o n s e - q u e n t e x p a n s i o n . o f i n f o r m a t i o n on b o t h c o n i f e r s and hardwoods (30, 31, 1 2 2 ) . As a r e s u l t , s u f f i c i e n t i n f o r m a t i o n i s a v a i l - a b l e on t h e p a t t e r n s o f t r a c h e i d l e n g t h v a r i a t i o n w i t h i n a t r e e and t h e p r o c e s s o f c e l l d i v i s i o n i n t h e c a m b i a l zone so t h a t some r e l a t i o n s h i p s a r e now q u i t e c l e a r . D e f i n i t e p a t t e r n s o f c e l l l e n g t h c h a n g es a r e e v i d e n t r a d i a l l y w i t h i n a g r o w t h r i n g . I t has b e e n shown by m a c e r a t i n g s e r i a l s e c t i o n s t h a t c e l l l e n g t h d e c r e a s e s f o l l o w i n g t h e i n - c e p t i o n o f y e a r l y g r o w t h u n t i l a minimum i s r e a c h e d e i t h e r i n t h e f i r s t p o r t i o n o f t h e e a r l y w o o d o r c l o s e t o t h e b o u n d a r y between t h e e a r l y w o o d and l a t e w o o d (15, 3 1 ) . T h e r e a f t e r , t h e c e l l l e n g t h i n c r e a s e s t o a maximum i n t h e l a t e w o o d , e i t h e r r a p i d l y o r s l o w l y , d e p e n d i n g upon t h e n a t u r e o f t r a n s i t i o n b etween t h e two i n t r a - i n c r e m e n t a l zones ( 1 5 ) . O t h e r s t u d i e s , - u s i n g i s o l a t e d e a r l y w o o d and l a t e w o o d m a t e r i a l f r o m t h e same i n c r e m e n t , h a v e c o n f i r m e d t h e r e s u l t s a l r e a d y c i t e d (31, 1 2 2 ) . Thus, l a t e w o o d t r a c h e i d s a r e on t h e a v e r a g e a b o u t 10 p e r c e n t l o n g e r t h a n e a r l y w o o d t r a c h e i d s . T r a c h e i d l e n g t h v a r i a t i o n v e r t i c a l l y w i t h i n an i n c r e m e n t f o l l o w s a g e n e r a l p a t t e r n o f g r a d u a l i n c r e a s e f r o m t h e b a s e upward t o a maximum below t h e crown, o r between 3 0 t o 4 0 p e r c e n t o f t h e t r e e h e i g h t (9,,30 13. 3 1 , 82, .98) and a d e c r e a s e i n l e n g t h a b o v e t h i s h e i g h t i n t h e t r u n k . R a d i a l v a r i a t i o n o f t r a c h e i d l e n g t h , i n s u c c e s s i v e g r o w t h i n c r e m e n t s , h a s b e e n i n v e s t i g a t e d i n a g r e a t number o f s t u d i e s , t h e r e s u l t s o f w h i c h a r e s u m m a r i z e d b e l o w . A t any g i v e n h e i g h t , t h e t r a c h e i d s o f - t h e f i r s t g r o w t h i n c r e m e n t a r e v e r y s h o r t ; c o n i f e r o u s t r a c h e i d s r a n g e i n l e n g t h f r o m 0.5 t o 1.5 mm ( 3 1 ) . L e n g t h i n c r e a s e s r a p i d l y i n t h e s e c o n d g r o w t h i n c r e m e n t a n d i n a few s u b s e q u e n t o n e s ; t h e n t h e r a t e o f i n c r e a s e d e c l i n e s , b u t c o n t i n u e s u n t i l a maximum l e n g t h i s r e a c h e d . I n c o n i f e r s t h i s may be t h r e e t o f i v e t i m e s g r e a t e r t h a n t h e i n i t i a l l e n g t h ( 2 6 , 3 1 ) . T h e r e i s no g e n e r a l r u l e as t o t h e number o f y e a r s n e c e s s a r y f o r r e a c h i n g maximum l e n g t h w h i c h v a r i e s w i t h t h e r a t e o f g r o w t h (137) . Maximum l e n g t h may b e r e a c h e d i n as l i t t l e as s i x t o e i g h t y e a r s , and may n o t be r e a c h e d u n t i l a t r e e i s 200 t o 300 y e a r s o f age; a c o n t i n u - ous i n c r e a s e h a s b e e n o b s e r v e d t h r o u g h o u t t h e g r o w t h i n c r e m e n t s o f a 4 5 5 - r y e a r - o l d D o u g l a s - ^ f i r t r e e (46) . A f t e r maximum i s r e a c h e d , t h e r e f o l l o w s a p e r i o d d u r i n g w h i c h l e n g t h may v a r y c o n s i d e r a b l y a b o u t a mean maximum l e n g t h ; f i n a l l y , i n v e r y o l d t r e e s , t h e c e l l l e n g t h may d e c r e a s e f r o m t h e maximum ( 5 ) . I n b o t h p a r t s o f t h e i n c r e m e n t , t r a c h e i d s h a v e a g e n e r a l i n c r e a s e i n l e n g t h f r o m p i t h t o b a r k , b u t e a r l y w o o d e x h i b i t s a more g r a d u a l r i s e i n t h e . e a r l y y e a r s t h a n i s shown by t h e l a t e w o o d ( 1 5 5 ) . F u r t h e r m o r e , t h e v a r i a b i l i t y i n l e n g t h f o r t h e e a r l y - . wood c e l l s i s much l e s s t h a n t h a t f o r l a t e w o o d c e l l s ( 3 0 , 1 5 5 ) . 14. T h e r e i s no g e n e r a l a g r e e m e n t w i t h r e s p e c t t o t h e r e l a t i o n s h i p o f i n c r e m e n t w i d t h t o c e l l l e n g t h , b u t t h e b a l a n c e o f o p i n i o n a p p e a r s t o be i n f a v o r o f a n e g a t i v e one; i . e . , w i t h i n a t r e e , w i d e r i n c r e m e n t s have s h o r t e r c e l l s o t h e r f a c t o r s s u c h as age b e i n g e q u a l (31, 40, 122, 1 3 7 ) . As r e g a r d s h e i g h t g r o w t h , a p o s i t i v e r e l a t i o n was. f o u n d t o e x i s t b etween t h i s and c e l l l e n g t h i n t h e f i r s t i n c r e m e n t f r o m p i t h ; t h u s . i n c o m p a r i n g two t r e e s o f d i f f e r e n t g r o w t h r a t e s , t h e t r e e o f more r a p i d h e i g h t g r o w t h was shown t o have t h e l o n g e r t r a c h e i d s (7, 23, 32, 82, 86, 1 0 6 ) . The p a t t e r n s o f v a r i a t i o n a r e o f t e n f o u n d t o be m o d i - f i e d i f t h e t r e e has u n d e r g o n e changes i n g r o w t h d i r e c t i o n f r o m v e r t i c a l , due t o e n v i r o n m e n t a l c h a n ges w h i c h i n v o l v e t h e f o r m a - t i o n o f s t r u c t u r a l l y m o d i f i e d t r a c h e i d , s u c h as t a k e s p l a c e d u r i n g t h e f o r m a t i o n o f c o m p r e s s i o n wood ( 1 4 5 ) , and c e l l l e n g t h d e c r e a s e s . T r a c h e i d l e n g t h between t r e e s i n a p a r t i c u l a r s p e c i e s i s a l s o i n f l u e n c e d by g e o g r a p h i c a l l o c a t i o n ( 8 ) . How- e v e r , i t i s t o be e m p h a s i z e d t h a t a l t h o u g h s l i g h t m o d i f i c a t i o n s o f t r a c h e i d l e n g t h c a n be i n d u c e d by e n v i r o n m e n t a l a g e n t s (108, 112, 1 1 3 ) , c o n s i d e r a b l e e v i d e n c e has b e en p r e s e n t e d f o r g e n e t i c c o n t r o l o v e r t r a c h e i d l e n g t h (8, 37, 47, 61, 100, 154, 1 6 9 ) . W i t h i n t h e t r e e , t r a c h e i d l e n g t h i s c o n t r o l l e d by p h y s i o l o g i c a l f a c t o r s . Thus D i n w o o d i e (32) c o n c l u d e d " t h a t t r a c h e i d l e n g t h v a r i a t i o n w i t h i n t h e t r e e i s a f u n c t i o n o f cambium w h i c h i s s t r o n g l y i n f l u e n c e d by t h e a c t i v i t y o f t h e t e r m i n a l m e r i s t e m , p r o b a b l y by means o f a u x i n . " S t u d i e s on t h e m e c h a n i s m s t h a t r e s u l t i n v a r i a t i o n s i n l e n g t h w i t h i n t h e t r e e a r e d i f f i c u l t , b u t t h e e x t e n s i v e i n v e s t i g a t i o n s o f B a n n a n (7, 8, 9) h a v e shown t h a t t h e r a t e o f a n t i c l i n a l d i v i s i o n s t a k i n g p l a c e i n t h e cambium i s a p r i n c i p a l f a c t o r . I t w o u l d be a p p r o p r i a t e t o q u o t e a r e c e n t s t a t e m e n t o f B a n n a n , i n t h i s r e g a r d ( 9 ) : I n g e n e r a l , r a t e o f a n t i c l i n a l d i v i s i o n a n d c e l l l e n g t h a r e n e g a t i v e l y r e l a t e d . T h i s a p p l i e s t o b o t h w i t h i n - t r e e v a r i a t i o n s a nd t o s p e c i e s - t o - s p e c i e s d i f f e r e n c e s . H o w e v e r , some" n o t a b l e e x c e p t i o n s o c c u r . Thus i n S e q u o i a s e m p e r v i r e n s , w h e r e t h e c e l l s a r e l o n g e r t h a n i n o t h e r c o n i f e r s , t h e r a t e o f ' a n t i c l i n a l d i v i s i o n i s s u b s t a n t i a l l y a b o v e a v e r a g e . . Con- v e r s e l y , P i n u s c o n t o r t a h a s b o t h r e l a t i v e l y s h o r t c e l l s a n d a l o w r a t e o f a n t i c l i n a l d i v i s i o n , an u n u s u a l c o m b i n a t i o n . I t t h e r e f o r e a p p e a r s t h a t t h e r a t e o f a n t i c l i n a l d i v i s i o n i n t h e cambium i s p r o b a b l y a p a r t o f t h e o p e r a t i v e m e c h a n i s m by w h i c h c e l l d i m e n s i o n s , d e t e r m i n e d by t h e i n t e r p l a y o f g e n e t i c , e n v i r o n m e n t a l a n d o n t o g e n e t i c f a c t o r s a r e r e g u l a t e d ( 9 ) . B a n n a n ' s w o r k i s s i g n i f i c a n t i n t h a t i t b r i d g e s e a r l i e r s t u d i e s o f c e l l s h a p e and d i m e n s i o n s w i t h p l a n t p h y s i o l o g i c a l i n v e s t i - g a t i o n s o f x y l e m d i f f e r e n t i a t i o n . D i m e n s i o n a l R e l a t i o n s h i p s 4 J n T r a c h e i d s I n v e s t i g a t i o n s o f v a r i a t i o n i n c e l l s i z e w i t h i n t h e t r e e w e r e p r i m a r i l y c o n c e r n e d w i t h c e l l l e n g t h . P o s s i b l e c h a n g e s o f c e l l d i a m e t e r and w a l l t h i c k n e s s w e r e g i v e n l i t t l e a t t e n t i o n . T h e s e and r e l a t e d a s p e c t s on c o n i f e r s w e r e d i s c u s s e d r e c e n t l y by L a r s o n ( 7 7 ) . Goggans ( 4 8 ) , w i t h s p e c i a l r e f e r e n c e t o s o u t h e r n p i n e s p e c i e s , r e v i e w e d t h e i m p o r t a n c e o f d i a m e t e r and w a l l t h i c k n e s s i n r e l a t i o n t o wood q u a l i t y . T a n g e n t i a l d i a m e t e r s a r e p r i m a r i l y d e t e r m i n e d by t h e w i d t h s o f c a m b i a l i n i t i a l s f r o m w h i c h c e l l s a r e d e r i v e d , and o r d i n a r i l y v e r y l i t t l e p o s t - d i v i s i o n e x p a n s i o n t a k e s p l a c e ( 2 ) . N e v e r t h e l e s s , t a n g e n t i a l t r a c h e i d d i a m e t e r s i n c r e a s e c o n s i d e r - a b l y as t h e w i d t h s o f c a m b i a l i n i t i a l s i n c r e a s e w i t h age on a c r o s s s e c t i o n ( 3 ) . As s t a t e d by L a r s o n ( 7 7 ) , " t h e , p r i n c i p a l v a r i a t i o n i n t r a c h e i d d i a m e t e r o c c u r s i n t h e r a d i a l d i r e c t i o n , and i t i s t h i s v a r i a t i o n t h a t c o n t r i b u t e s t o l a t e w o o d f o r m a t i o n and t h e s t r u c t u r a l d i f f e r e n c e s i n t h e wood w i t h i n and between t r e e s . " A g e n e r a l c o n c l u s i o n o f Mohr and R o t h (95) was t h a t t h e r a d i a l w i d t h o f t r a c h e i d s i n c r e a s e s f r o m t h e p i t h o u t w a r d and t h a t t h e i n c r e a s e was r a p i d d u r i n g t h e e a r l y y e a r s . L a r s o n (77) q u o t e d s e v e r a l a u t h o r s and c o n c l u d e d t h a t , w i t h i n c r e a s i n g age, r a d i a l d i a m e t e r g e n e r a l l y t e n d s t o i n c r e a s e t o a maximum w i d t h and f l u c t u a t e s t h e r e a f t e r . T h i s i s p r i m a r i l y a p p l i c a b l e t o e a r l y - wood t r a c h e i d s s i n c e no d e f i n i t e age p a t t e r n c a n be e s t a b l i s h e d f o r l a t e w o o d t r a c h e i d s . W i t h i n a s i n g l e g r o w t h i n c r e m e n t , e a r l y w o o d t r a c h e i d d i a m e t e r u s u a l l y i n c r e a s e s f r o m t h e stem b a s e t o a maximum a t some p o i n t i n mid-stem, t h e n d e c r e a s e s t o w a r d s t h e apex. 17. I n v e s t i g a t i o n s o f L a r s o n ( 7 7 ) , R i c h a r d s o n ( 1 1 2 ) a n d W o d z i c k i ( 1 6 4 ) i n d i c a t e t h a t t h e p h y s i o l o g i c a l p r o c e s s e s c o n t r o l l i n g s e c o n d a r y w a l l t h i c k n e s s a r e i n d e p e n d e n t o f t h o s e r e g u l a t i n g t r a c h e i d d i a m e t e r , a l t h o u g h t h e s e two c h a r a c t e r i s t i c s do v a r y more o r l e s s s i m u l t a n e o u s l y - i n n a t u r e . W hereas t r a c h e i d d i a m e t e r was p r i m a r i l y d e t e r m i n e d by t h e amount o f a u x i n r e a c h i n g a d e v e l - o p i n g t r a c h e i d , w a l l t h i c k n e s s a p p e a r s t o be d e t e r m i n e d by t h e amount o f s u c r o s e , o r p h o t o s y n t h a t e , r e a c h i n g e a c h t r a c h e i d ( 8 0 ) . T r a c h e i d w a l l t h i c k n e s s i n c r e a s e s w i t h age on a c r o s s s e c t i o n t o a maximum t h a t v a r i e s w i t h g r o w t h c o n d i t i o n s ( 6 5 , 77, 1 2 0 ) . W i t h i n a g r o w t h i n c r e m e n t , w a l l t h i c k n e s s d e c r e a s e s w i t h h e i g h t t o some p o i n t i n m i d - s t e m and may e x h i b i t a s l i g h t i n c r e a s e w i t h i n t h e c r o w n . T h e s e p a t t e r n s r e f e r m o s t l y t o l a t e w o o d t r a c h e i d s a n d , a l t h o u g h e a r l y w o o d t r a c h e i d w a l l t h i c k n e s s a l s o v a r i e s , t h e v a r i a t i o n s a r e m i n i m a l ( 7 7 ) . H i l l e r (57) s t u d i e d v a r i a t i o n i n r a d i a l w a l l t h i c k n e s s o f l a t e w o o d c e l l s i n s e v e r a l e n t i r e a n n u a l g r o w t h s h e a t h s o f l o b l o l l y p i n e ( P i n u s t a e d a L . ) . She f o u n d t h a t w a l l t h i c k n e s s i n c r e a s e d r a p i d l y f r o m t h e s e c o n d t o a b o u t t h e e i g h t h i n t e r n o d e f r o m t h e s h e a t h a p e x , t h e n more s l o w l y t o t h e t r e e b a s e . L a r s o n (78) showed t h a t i n r e d p i n e ( P i n u s r e s i n o s a A i t . ) e a r l y w o o d w a l l t h i c k n e s s i n c r e a s e d g r a d u a l l y w i t h a g e , w h e r e a s i n l a t e w o o d t h e i n c r e a s e was more r a p i d b u t e x h i b i t e d g r e a t e r f l u c t u a t i o n s . F i n a l l y , t h e c e l l s t h a t c o m p r i s e t h e o u t e r g r o w t h i n c r e m e n t o f v e r y o l d t r e e s (320 t o 400 y e a r s ) a r e o f t e n t h i n w a l l e d i n com- . p a r i s o n w i t h wood f o r m e d d u r i n g e a r l i e r s t a g e s o f g r o w t h ( 5 2 ) . 18. W i t h i n a t r e e there i s some evidence t h a t the micro- f i b r i l o r i e n t a t i o n , and to l e s s e r degree the f i b r e composition, i s r e l a t e d to the f i b r e dimension (145). A r e l a t i o n s h i p between m i c r o f i b r i l o r i e n t a t i o n i n the c e l l w a l l and the morphology of wood f i b r e s was f i r s t r e p o r t e d as long ago as 1934 by P r e s t o n (109, 110). He observed t h a t f o r the l a y e r i n gymnosperms the angle of o r i e n t a t i o n of the c e l l u l o s e molecules (micro- f i b r i l s ) was g r e a t e r i n the s h o r t f i b r e s of e a r l y growth i n c r e - ments than i n longer f i b r e s of l a t e r ones. The r e l a t i o n s h i p was shown to apply to the t r a c h e i d s from s u c c e s s i v e growth increments as w e l l as to random samples from a number of growth increments, so t h a t the r e l a t i o n s h i p d i d not r e f l e c t a process of aging of meristem from which the t r a c h e i d s were d e r i v e d (see a l s o 142, 143, 144, 145). T h i s was subsequently shown to apply f o r the (145) and f o r the l a y e r s (89), as w e l l as i n the abnormal t r a c h e i d s of compression wood (145, 146), f i b r e s of angiosperms (145), phloem f i b r e s (110) and c o t t o n f i b r e s (145). I t i s of i n t e r e s t to note t h a t H i l l e r (57) s i m i l a r l y found a high degree of c o r r e l a t i o n between r a d i a l w a l l t h i c k n e s s and f i b r i l angle i n t h a t , w i t h i n c r e a s e i n w a l l t h i c k n e s s , t h e r e i s decrease i n f i b r i l angle. L i k e w i s e , Necesany (97) showed t h a t m i c r o f i b r i l t h i c k n e s s , c e l l w a l l t h i c k n e s s , and c e l l l e n g t h are f e a t u r e s of mutual d i r e c t p r o p o r t i o n a l i t y . He f u r t h e r noted t h a t an i n v e r s e p r o p o r t i o n a l r e l a t i o n s h i p e x i s t s between m i c r o f i b r i l t h i c k n e s s and l i g n i n content i n the f i b r e . T h e r e a r e i n d i c a t i o n s t h a t c e l l l e n g t h and w i d t h a r e a l s o r e l a t e d . Thus a number o f i n v e s t i g a t o r s , among them Bannan (6) , G r a f f and M i l l e r (50) , H a r l o w (54) , H e i n i g r and Simmonds ( 5 6 ) , and W h e e l e r e t a l . ( 1 5 8 ) , have d e m o n s t r a t e d t h a t t h e r e i s a d i r e c t p o s i t i v e r e l a t i o n s h i p between f i b r e l e n g t h and d i a m e t e r w i t h i n e a c h s p e c i e s . I n a d d i t i o n , a r e c e n t s t u d y by V o r r e i t e r (140) b r o u g h t o u t a r e c t i l i n e a r i n c r e a s e i n f i b r e l e n g t h w i t h i n c r e a s i n g t h i c k n e s s o f f i b r e w a l l , f o r c o n i f e r o u s s p e c i e s . . R e s u l t s o f P r e s t o n (109, 110) a n d - H i l l e r , ( 5 7 ) , i f c o n s i d e r e d t o g e t h e r , a l s o p o i n t o u t t h a t w a l l t h i c k n e s s and c e l l l e n g t h may be r e l a t e d s i n c e b o t h t h e s e f e a t u r e s a r e i n - v e r s e l y c o r r e l a t e d w i t h m i c r o f i b r i l a n g l e . The r e v i e w p r e s e n t e d above shows t h a t d e f i n i t e i n t e r - r e l a t i o n s h i p s e x i s t between c e l l d i m e n s i o n s , i n p a r t i c u l a r l e n g t h , and w a l l o r g a n i z a t i o n ; h e n c e w i t h t r a c h e i d c o m p o s i t i o n . C h e m i c a l C o m p o s i t i o n and R e c o v e r y o f C a r b o h y d r a t e F r a c t i o n Wood c o n s i s t s o f t h r e e m a i n c o n s t i t u e n t s , namely c e l l u l o s e , h e m i c e l l u l o s e s , and l i g n i n ( 1 3 5 ) . H o l o c e l l u l o s e i s t h e m a j o r c o n s t i t u e n t , and i s , by d e f i n i t i o n , t h e l i g n i n - f r e e f i b r o u s m a t e r i a l c o m p r i s i n g a l l o f t h e h e m i c e l l u l o s e s and c e l l u - l o s e i n wood ( 1 2 8 ) . The t o t a l c e l l w a l l p o l y s a c c h a r i d e s a r e u s u a l l y o b t a i n e d f r o m wood s u b s t a n c e i n t h e f o r m o f h o l o c e l l u - lose/,of which a n g i o s p e r m s c o n t a i n f r o m 70 t o 80 p e r c e n t , and 20. c o n i f e r s 60 t o 75 p e r c e n t , b a s e d on e x t r a c t i v e - f r e e wood. The c h i e f v a l u e o f h o l o c e l l u l o s e p r e p a r a t i o n i s t h a t i t o f f e r s a b e g i n n i n g m a t e r i a l f o r f u r t h e r r e s e a r c h . G l u c a n , as a p p r o x i m a t e d by a l p h a - c e l l u l o s e , c o n s t i t u t e s 45 t o 50 p e r c e n t o f h o l o c e l l u l o s e i n a n g i o s p e r m s and 40 t o 45 p e r c e n t o f t h a t i n c o n i f e r s ( 2 0 ) ; t h i s i s c u s t o m a r i l y r e g a r d e d as t h e t r u e c e l l u l o s i c f r a c t i o n o f wood c a r b o h y d r a t e s . A l p h a - c e l l u l o s e i s c o n s i d e r e d t o be t h a t f r a c t i o n o f h o l o c e l l u l o s e o r p u l p w h i c h i s i n s o l u b l e i n a s o l u - t i o n o f 17.5 p e r c e n t NaOH a t 20°C ( 1 3 1 ) . The t e r m , h o w e v e r , i s s t r i c t l y a r b i t r a r y , and d o e s n o t i m p l y e x c l u s i v e l y a homogeneous g l u c a n f o r , i n v a r i a b l y , i t c o n t a i n s s m a l l amounts o f c a r b o n d i o x i d e - and f u r f u r a l - y i e l d i n g m a t e r i a l s , as w e l l as s i g n i f i c a n t amounts o f mannans f r o m c o n i f e r s ( 1 2 6 , 1 6 2 , 1 6 3 ) . S t a n d a r d methods f o r d e t e r m i n i n g h o l o c e l l u l o s e o r s i m i l a r c a r b o h y d r a t e f r a c t i o n s , r e q u i r i n g 2 g o f wood m e a l p e r s a m p l e , a r e a v a i l a b l e ( 1 2 8 , 1 2 9 ) . I n a d d i t i o n , f o u r m i c r o - m e t h o d s , r e q u i r i n g a b o u t 0.5 g s a m p l e s , h a v e b e e n a d v a n c e d f o r d e t e r m i n a t i o n o f h o l o c e l l u l o s e o r s i m i l a r c a r b o h y d r a t e f r a c t i o n ( s e e R e f . 1 6 1 ) . Z o b e l and M c E l w e e (171) and E r i c k s o n (43) a p p l i e d c h l o r i t e , W a t s o n (149) u s e d c h l o r i n e - s u l p h a t e , and L e o p o l d (83) e m p l o y e d p e r a c e t i c a c i d f o l l o w e d by s o d i u m b o r o - h y d r i d e r e d u c t i o n . A f i f t h s e m i - m i c r o - m e t h o d has b e e n u s e d by M e i e r (90 , 91) and L a r s o n (78) i n w h i c h s e c o n d a r y x y l e m f i b r e s a r e h y d r o l y z e d t o t h e i r m o n o s a c c h a r i d e s , f r o m w h i c h t h e p o l y - s a c c h a r i d e c o m p o s i t i o n i s c a l c u l a t e d f r o m some b a s i c a s s u m p t i o n s 21. and c h r o m a t o g r a p h i c e v i d e n c e . A c o m p r e h e n s i v e r e v i e w o f l i t e r a t u r e on t h e s e a s p e c t s i s c o n t a i n e d i n a t h e s i s by S q u i r e ( 1 2 3 ) . S q u i r e a l s o d e v e l o p e d a new m i c r o - m e t h o d f o r c e l l u l o s e d e t e r m i n a t i o n , and q u a n t i t a t i v e l y e s t i m a t e d i n t r a - i n c r e m e n t a l a l p h a - c e l l u l o s e y i e l d as t h e c o r r e c t e d y i e l d o f n i t r a t e d wood m e a l . A m a j o r l i m i t a t i o n o f h i s t e c h n i q u e was t h a t i t - c o u l d n o t be a p p l i e d t o a l l woods. D i f f e r e n c e s i n a n a l y t i c a l p r o c e d u r e s w i l l show v a r i a - t i o n o f c h e m i c a l c o m p o s i t i o n i n t h e same sample o f wood ( 1 3 7 ) . F o r example, t h e C r o s s and Bevan c e l l u l o s e (128) does n o t c o n - t a i n t h e e n t i r e h e m i c e l l u l o s e f r a c t i o n as some i s e x t r a c t e d w i t h t h e l i g n i n . Thus t h e p u b l i s h e d d a t a on h o l o c e l l u l o s e f o r a p a r t i c u l a r s p e c i e s o f wood a r e i n c o n s i s t e n t . As s u c h , t h e most r i g o r o u s e v a l u a t i o n o f a n a l y t i c a l d a t a i s r e p r e s e n t e d by summative a n a l y s e s , i n w h i c h t h e a n a l y s t aims t o a c c o u n t f o r a l l c o n s t i t u e n t s p r e s e n t by a summation t h a t i d e a l l y s h o u l d t o t a l 100 p e r c e n t , i f no c o n s t i t u e n t s have b e e n o v e r l o o k e d and t h e r e i s no o v e r l a p p i n g ( 2 0 ) . A summation t h a t i n c l u d e s e x t r a c - t i v e s , l i g n i n , h o l o c e l l u l o s e , and a s h i d e a l l y a c c o u n t s f o r a l l c h e m i c a l - c o n s t i t u e n t s o f wood. Many w o r k e r s p r e f e r , however, t o u s e p r e - e x t r a c t e d wood and t o r e p o r t summative a n a l y s e s on t h e e x t r a c t i v e - f r e e and m o i s t u r e - f r e e b a s i s , b e c a u s e o f p o s s i b l e e r r o r s t h a t may a r i s e f r o m v a r i a b i l i t y i n e x t r a c t i v e c o n t e n t i n d i f f e r e n t woods ( 2 0 ) . I n c o n i f e r s , t h e p r o p o r t i o n s o f c e l l u l o s e , h e m i c e l l u - l o s e s , and l i g n i n , b a s e d on s t a n d a r d i z e d methods o f p r e p a r a t i o n , a r e a p p r o x i m a t e l y as f o l l o w s ( p e r c e n t o f e x t r a c t i v e - f r e e , d r y wood): c e l l u l o s e 40 t o 45 p e r c e n t , l i g n i n 25 t o 35 p e r c e n t , and h e m i c e l l u l o s e s 20 f o . 2 5 p e r - c e n t (20', 135). v The p r o p o r t i o n o f p e c t i c s u b s t a n c e s i s s m a l l ( a b o u t 1 p e r c e n t ) ( 1 3 5 ) . I n t e r - s p e c i e s v a r i a t i o n i n a l l c h e m i c a l components, f o r s e v e r a l c o n i - f e r s , i s c o n v e n i e n t l y l i s t e d by B r o w n i n g (20), T i m e l l (135, 136) and S q u i r e ( 1 2 3 ) . B r o w n i n g (20) a l s o has shown wood e l e m e n t a l a n a l y s e s f o r some o f t h e s e s p e c i e s . G e n e r a l l y , c h e m i c a l v a r i a - t i o n b etween s p e c i e s i s l e s s t h a n t h a t b etween g e n e r a ; however, many e x c e p t i o n s h a v e b e e n r e p o r t e d . The c h e m i c a l c o m p o s i t i o n o f r e a c t i o n wood d i f f e r s f r o m t h a t o f n o r m a l wood i n s e v e r a l r e s p e c t s (136) . T e n s i o n wood c o n t a i n s l e s s l i g n i n and x y l a n b u t more c e l l u l o s e and more g a l a c t o s e r e s i d u e s t h a n n o r m a l wood. C o m p r e s s i o n wood has a h i g h l i g n i n and a low c e l l u l o s e c o n t e n t compared t o n o r m a l wood. The r e l a t i v e amount o f mannose r e s i d u e i s much r e d u c e d . The p r e d o m i n a n t h e m i c e l l u l o s e i s i n s t e a d a p o l y s a c c h a r i d e b a s e d on g a l a c t o s e r e s i d u e s . P i l l o w and Bray- (107) r e p o r t e d t h a t l o b l o l l y p i n e a l p h a - c e l l u l o s e , f r o m C r o s s and Bevan a n a l y s i s , v a r i e d f r o m 46 p e r c e n t i n n o r m a l wood t o 3 5 p e r c e n t i n c o m p r e s s i o n wood. L i g n i n c o n t e n t i n c r e a s e d f r o m 28 t o 3 5 p e r c e n t , r e s p e c - t i v e l y , w i t h no s i g n i f i c a n t change i n t o t a l p e n t o s a n s . A p p a r - e n t l y , t h e i n c r e a s e i n l i g n i n o c c u r s a t t h e e x p e n s e o f c e l l u l o s e a n d / o r g l ucomannan f o r c o n i f e r s , and t h e r e v e r s e seems t o o p e r a t e i n t h e . r e a c t i o n wood c h e m i s t r y o f hardwoods. T h e r e i s some e v i d e n c e t h a t c o m p r e s s i o n wood a l s o d i f f e r s f r o m n o r m a l wood, q u a l i t a t i v e l y . T h u s , D i n w o o d i e ' s work (33) i n d i c a t e s low c e l l u l o s e DP ( d e g r e e o f p o l y m e r i z a t i o n ) f o r c o m p r e s s i o n wood f i b r e s . An e x t e n s i v e r e v i e w o f l i t e r a t u r e on a n a t o m i c a l and c h e m i c a l p r o p e r t i e s o f c o m p r e s s i o n wood i s g i v e n by W e s t i n g (151, 1 5 2 ) . A n o t h e r a s p e c t r e l a t e d t o t h e t h e s i s t o p i c i n c l u d e s v a r i a t i o n s i n c a r b o h y d r a t e c o n t e n t a c r o s s a t r e e d i a m e t e r , w h i c h a r e s i g n i f i c a n t b e c a u s e o f j u v e n i l e wood. Z o b e l and McElwee (171) and B y r d e t a l . (22)^ r e p o r t e d t h a t , b a s e d on d r y w e i g h t , l o b l o l l y p i n e o u t e r wood c o n t a i n e d one t o t h r e e p e r c e n t more h o l o c e l l u l o s e , and f o u r t o s e v e n p e r c e n t more a l p h a - c e l l u l o s e t h a n t h a t o f j u v e n i l e wood f r o m the.same t r e e s . I n g e n e r a l , i t a p p e a r s t h a t , f o r c o n i f e r o u s wood, t h e r e i s a r a p i d i n i t i a l i n c r e a s e i n c e l l u l o s e c o n t e n t f r o m t h e i n c r e m e n t s c l o s e t o t h e p i t h , and somewhat l e s s change b e y o n d t h e j u v e n i l e c o r e . I t was shown by Kennedy and J a w o r s k y ( 6 9 ) , f o r D o u g l a s - f i r , t h a t C r o s s and Bevan c e l l u l o s e y i e l d s i n c r e a s e d f r o m 57.4 p e r c e n t i n i n c r e m e n t s 1 t o 5 t o 61.6 p e r c e n t i n i n c r e m e n t s 16 t o 25. O n l y a modest f u r t h e r i n c r e a s e o f 1.7 p e r c e n t was r e p o r t e d i n t h e r e m a i n d e r o f t h e 8 0 - y e a r - o l d t r e e . L i k e w i s e , Wardrop (141) o b s e r v e d an i n c r e a s e i n C r o s s and Bevan c e l l u l o s e f r o m 58 t o 65 p e r c e n t w i t h i n t h e f i r s t 20 i n c r e m e n t s f r o m p i t h , f o r t h e same s p e c i e s . C h l o r i t e h o l o c e l l u l o s e and a l p h a - c e l l u l o s e p a t t e r n s a l s o f o l l o w t h e above t r e n d . H o l o c e l l u l o s e p e r c e n t a g e s a p p e a r t o i n c r e a s e l e s s r a p i d l y t h a n a l p h a - c e l l u l o s e (52, 6 9 ) , l e a d i n g t o t h e c o n c l u s i o n t h a t t h e r e may be a g r e a t e r h e m i c e l l u l o s e f r a c t i o n i n i n c r e m e n t s c l o s e t o t h e p i t h ( 7 0 ) . T h i s may be. a t t r i b u t e d p a r t l y t o t h e a u t o h y d r o l y s i s o f t h e o l d e r c e l l w a l l m a t e r i a l c l o s e r t o t h e p i t h ( 7 0 ) . A p o i n t o f i n t e r e s t i s t h e s i m i l a r i t y i n b e h a v i o u r o f g r o s s wood c e l l u l o s e c o n t e n t i n s u c c e s s i v e i n c r e m e n t s f r o m t h e p i t h o u t w a r d , and t h a t o f t r a c h e i d l e n g t h a c r o s s t h e stem. T h i s s i m i l a r i t y was shown by H a l e and C l e r m o n t (52) w o r k i n g w i t h D o u g l a s - f i r , and by Wardrop ( 1 4 1 ) , w i t h t h e same s p e c i e s and w i t h r a d i a t a p i n e ( P i n u s r a d i a t a (D. Don.). H a l e and C l e r m o n t a l s o r e p o r t e d a low a l p h a - c e l l u l o s e p e r c e n t a g e i n o v e r m a t u r e wood (322 t o 400 y e a r s ) t h a t g e n e r a l l y c o n t a i n e d t h i n - w a l l e d p r o s e n c h y m a c e l l s . They a t t r i b u t e d t h i s t o t h e e x t r e m e l y s l o w g r o w t h o f wood a t t h a t a ge. O t h e r f a c t o r s t h a t c o u l d c o n t r i b u t e t o t h e d i f f e r e n c e s i n c a r b o h y d r a t e amount i n v o l v e v a r i a t i o n w i t h i n t h e g r o w t h i n c r e m e n t . Q u a n t i t a t i v e d i f f e r e n c e s i n g r o w t h zone c h e m i s t r y were f i r s t r e p o r t e d i n 1926 by R i t t e r and F l e c k ( 1 1 4 ) . They o b s e r v e d t h a t c e l l u l o s e forms a l a r g e r p e r c e n t a g e (up t o 3.4 p e r c e n t ) o f t h e t o t a l wood s u b s t a n c e i n l a t e w o o d t h a n i n e a r l y w o o d , o f t h r e e c o n i f e r o u s woods, and an o p p o s i t e s i t u a t i o n p r e v a i l s f o r l i g n i n . T h e s e r e s u l t s were c o n f i r m e d f o r c o n i f e r s by s u b s e q u e n t w o r k e r s f o r b o t h h o l o - and a l p h a - c e l l u l o s e f r a c t i o n s (52, 123, 124, 150, 1 6 1 ) . Some o f t h e l a t e r w o r k e r s , however, showed g r e a t e r d i f f e r e n c e i n t h e m a g n i t u d e o f y i e l d s t h a n r e p o r t e d by R i t t e r and F l e c k . An anomaly o c c u r s h e r e , i n / t h a t t h e f i r s t - f o r m e d e a r l y w o o d a p p e a r s t o r e t a i n some s i m i l a r i t y a t t h e c h e m i c a l l e v e l o f o r g a n i z a t i o n t o l a s t - f o r m e d t i s s u e s o f t h e p r e c e d i n g s e a s o n . L a t e r - f o r m e d e a r l y w o o d ( i . e . , f r o m t h e p r e s e n t y e a r ) , a c c o r d i n g t o S q u i r e ( 1 2 3 ) , does n o t seem t o r e t a i n s u c h s i m i - l a r i t y . T h u s , minimum e s t i m a t e d c e l l u l o s e y i e l d o c c u r r e d a t c o n s i d e r a b l e r a d i a l - c e l l u l a r • depth 1. F u r t h e r a n a l y s e s a t - t h e i n t r a - i n c r e m e n t a l . l e w e l ; r e p o r t e d by-Wellwood. and .Wilson (156) , i n d i c a t e t h a t peak c h l o r i t e h o l o c e l l u l o s e (77 t o 78 p e r c e n t , c o r r e c t e d f o r l i g n i n ) o c c u r s a t t h e p o i n t o f l a t e w o o d i n i t i a t i o n , w h i l e t h e f i r s t - and l a s t - f o r m e d c e l l s o f i n c r e m e n t s had l o w e r v a l u e s (72.5 t o 73.5 p e r c e n t ) , i n t h r e e a d j a c e n t i n c r e m e n t s o f D o u g l a s - f i r . B e s i d e s q u a n t i t a t i v e d i f f e r e n c e s > q u a l i t a t i v e d i f f e r e n c e s between e a r l y w o o d and l a t e w o o d were n o t e d . I t was shown t h a t , f o r D o u g l a s - f i r , t h e a v e r a g e c e l l u l o s e c h a i n l e n g t h o f l a t e w o o d i s s i x p e r c e n t h i g h e r t h a n t h a t o f e a r l y w o o d ( 1 6 1 ) . C a r b o h y d r a t e c o n t e n t i n t h e wood c o u l d be i n f l u e n c e d by s i t e i n d e x and t h e a s s o c i a t e d t r e e p r o p e r t i e s s u c h as g r o w t h r a t e , d i a m e t e r a t b r e a s t h e i g h t , t o t a l h e i g h t , and s p e c i f i c g r a v i t y o f t h e wood. Th u s , i t was shown by W o r s t e r and Sugiyama (165) t h a t s i t e i n d e x , w h i c h i s a c o m p o s i t e . o f s e v e r a l b a s i c f a c t o r s , h as a p r o n o u n c e d i n f l u e n c e on c a r b o h y d r a t e c o n t e n t o f c o n i f e r s and a n g i o s p e r m s . C o n t r a r y t o t h i s , M c M i l l a n (87) f o u n d t h a t t h e w i d t h o f a n n u a l i n c r e m e n t does n o t i n f l u e n c e p o l y - s a c c h a r i d e c o n t e n t when s p e c i f i c g r a v i t y e f f e c t s a r e e l i m i n a - t e d . M e i e r ' s (90) a n a l y s i s c o n f i r m s t h i s c o n c e p t ; he n o t e s t h a t much o f t h e v a r i a t i o n i n p o l y s a c c h a r i d e c o n c e n t r a t i o n i n t h e d i f f e r e n t c e l l w a l l l a y e r s i s r e l a t e d t o d e n s i t y . T h e r e - f o r e , t h e o b s e r v e d d i f f e r e n c e s i n e a r l y w o o d and l a t e w o o d c a r - b o h y d r a t e c o n t e n t s c o u l d be r e l a t e d t o s p e c i f i c g r a v i t y o r amount o f c e l l w a l l s u b s t a n c e p e r u n i t volume i n t h e wood (124) . E s t i m a t e s o f C a r b o h y d r a t e F r a c t i o n and W e i g h t i n R e l a t i o n t o C e l l u l a r E l e m e n t s I n t e r e s t i n t h e i n t e n s i v e e x a m i n a t i o n o f i n d i v i d u a l c e l l s i n woody t i s s u e has c e n t r e d m o s t l y on d e t e r m i n i n g t h e w a l l p h y s i c a l o r g a n i z a t i o n , and d i s t r i b u t i o n o f c h e m i c a l c o n - s t i t u e n t s i n t h e w a l l l a y e r s . The w e i g h t f a c t o r (amount o f c a r b o h y d r a t e f r a c t i o n ) o f i n d i v i d u a l c e l l s and i t s r e l a t i o n t o any p a r t i c u l a r c e l l e l e m e n t has been s t u d i e d v e r y l i t t l e . Some s t u d i e s (10, 104, 105) r e p o r t e d t h e p e r c e n t a g e s ( w e i g h t b a s i s ) o f v e r t i c a l t r a c h e i d s (98 t o 99 p e r c e n t ) and r a y parenchyma (one t o two p e r c e n t ) by q u a n t i t a t i v e f r a c t i o n a t i o n o f t h e t o t a l h o l o c e l l u l o s e d e r i v e d f r o m wood. I n o n l y one i n s t a n c e h a v e c o m p o s i t i o n a l d a t a b e e n r e l a t e d t o t h e w e i g h t o f a b i o l o g i c a l u n i t . T h u s , T h o r n b e r and N o r t h c o t e (13 4) have e s t i m a t e d t h e a v e r a g e w e i g h t and l e n g t h o f s i n g l e c e l l s i n f o u r s p e c i e s c o n s i s t i n g o f t h r e e a n g i o s p e r m s and one c o n i f e r . They f o u n d t h a t t h e mean w e i g h t o f t h e c e l l i n c r e a s e d w i t h age by as much as 3 50 t o 930 p e r c e n t i n t h e f o u r s p e c i e s examined by them. I t was c o n c l u d e d t h a t , a t a l l s t a g e s , t h e c e l l s f r o m t h e c o n i - f e r w e r e . c o n s i d e r a b l y h e a v i e r t h a n t h o s e f r o m t h e a n g i o s p e r m s . No a t t e m p t was made t o r e l a t e t h e o b s e r v e d w e i g h t d i f f e r e n c e s i n c e l l s t o l e n g t h . . E s t i m a t e s o f t r a c h e i d w e i g h t s were made by Y i a n n o s (166) i n c o n n e c t i o n w i t h c e l l w a l l d e n s i t y o f c o m m e r c i a l p u l p f i b r e s . W e i g h t s were b a s e d on measurement o f a g r o u p o f f i b r e s u s i n g a M e t t l e r UM7 b a l a n c e , and e x p r e s s e d as w e i g h t p e r u n i t l e n g t h ( c o a r s e n e s s ) . An i n g e n i o u s method o f w e i g h i n g s i n g l e t r a c h e i d s was f i r s t a t t e m p t e d by Skene (121) u s i n g a q u a r t z f i l a m e n t b a l a n c e . W e i g h t s r e p o r t e d r a n g e d f r o m 0.04 t o 0.1 y g . N e i t h e r t h e s p e c i e s e m ployed, n o r t h e t r a c h e i d c o m p o s i t i o n a r e a v a i l a b l e . W e i g h i n g c e l l s and r e l a t i n g r e s u l t s t o a m o r p h o l o g i c a l p r o p e r t y has s e v e r a l a d v a n t a g e s o v e r m e a s u r i n g l a t e r a l dimen- s i o n s . I t i s n o t o n l y much e a s i e r and q u i c k e r , b u t i n c l u d e s a l s o t h e e f f e c t s o f c e l l w a l l t h i c k n e s s , t h e s i z e o f t h e lumen,and the" amount o f c e l l u l o s i c m a t e r i a l c o m p o s i n g t h e c e l l s . T h i s p r o p e r t y , i . e . , f i b r e w e i g h t , i s u s u a l l y d e s c r i b e d as ' c o a r s e - n e s s ' and i s d e f i n e d as t h e w e i g h t p e r u n i t l e n g t h ; i t i s e x p r e s s e d i n m i l l i g r a m s p e r 100 m and c a l l e d a d e c i g r e x , a b b r e v i a t e d t o 'dg' ( 1 3 2 ) . I n v i e w o f i t s i m p o r t a n t e f f e c t on 28. many p r o p e r t i e s o f p a p e r , t h e . c o n c e p t o f f i b r e ' c o a r s e n e s s ' (24) and a method f o r i t s measurement f o r p u l p s a m p l e s , h a ve b e e n d e s c r i b e d i n TAPPI S u g g e s t e d Method T234 su-67 ( 1 3 2 ) . A new method f o r d e t e r m i n a t i o n o f f i b r e c o a s r e n e s s i n wood, b a s e d upon t h e number o f f i b r e s p e r s q u a r e mm o f c r o s s s e c t i o n and t h e s p e c i f i c g r a v i t y , was d e s c r i b e d by B r i t t (16, 1 7 ) . F i b r e c o a r s e n e s s measurements on v a r i o u s w e s t e r n c o n i - f e r o u s s p e c i e s , a t v a r i o u s a g e s , were made f o l l o w i n g t h i s t e c h - n i q u e ( 1 6 ) . I t was shown f r o m s u c h s t u d i e s t h a t t h e r e was a p r o g r e s s i v e b u t somewhat i r r e g u l a r i n c r e a s e i n f i b r e c o a r s e - n e s s w i t h i n c r e a s i n g age o f t h e t r e e , i n d i c a t i n g t h a t , p e r u n i t l e n g t h , a f i b r e f o r m e d as t h e t r e e grows o l d e r w e i g h s more. I t i s a l s o known t h a t f i b r e l e n g t h i n c r e a s e s f r o m c e n t r e o f a t r e e o u t w a r d ; t h u s • C l a r k \ ( 2 4 ) , b a s e d on s t u d i e s on p u l p s a m p l e s has shown t h a t p e r u n i t l e n g t h , l o n g e r f i b r e s w e i g h more t h a n s h o r t e r o n e s . S u c h a c o n c l u s i o n c a n be j u s t i f i e d on t h e b a s i s o f known i n t e r - r e l a t i o n s h i p s w i t h i n t h e t r a c h e i d m o r p h o l o g i c a l p r o p e r t i e s d i s c u s s e d e a r l i e r . N o t o n l y t h e w e i g h t s , b u t c h e m i c a l c o m p o s i t i o n o f s p e c i f i c wood c e l l s d i f f e r (1, 55, 103, 133, 1 4 2 ) . The h i g h e r p e n t o s a n c o n t e n t o f r a y c e l l s compared t o t h e e n t i r e wood has b e e n known f o r some t i m e ( 1 , 55, 133, 1 3 6 ) . R e c e n t s t u d i e s o f P e r i l a and h i s c o - w o r k e r s (103, 104, 105) c o n f i r m t h i s f o r b o t h a n g i o s p e r m s and c o n i f e r s . T h e i r d a t a i n d i c a t e t h a t t h e c h a r a c - t e r i s t i c c h e m i c a l f e a t u r e o f t h e parenchyma c e l l s , as compared t o p r o s e n c h y m a , i s t h e i r u n u s u a l l y h i g h x y l a n c o n t e n t . E f f e c t s o f F e r t i l i z a t i o n on Wood P r o p e r t i e s and C h e m i c a l C o m p o s i t i o n Klem (71) , i n a r e c e n t p a p e r , summarized t h e m a i n e f f e c t s o f f e r t i l i z a t i o n o f f o r e s t s t a n d s on wood c h a r a c t e r i s t i c s o f s p r u c e and p i n e s p e c i e s . P r o b l e m s i n a s s e s s i n g wood q u a l i t y o f f e r t i l i z e d c o n i f e r o u s t r e e s were d i s c u s s e d by L a r s o n ( 7 9 ) . D e t a i l e d l i t e r a t u r e s u r v e y s h a v e a l s o b e en p r o v i d e d by L e e ( 8 1 ) , P o s e y ( 1 0 8 ) a n d S a s t r y ( 1 1 7 ) . C o n s e q u e n t l y , s u c h i n f o r m a t i o n i s n o t p r e s e n t e d i n d e t a i l i n t h i s t h e s i s . I n most i n s t a n c e s , t h e a p p l i c a t i o n o f f e r t i l i z e r s t o f o r e s t s t a n d s r e s u l t s i n an i n c r e a s e d g r o w t h r e s p o n s e and a s l i g h t change i n t h e e a r l y w o o d - l a t e w o o d r a t i o . Changes i n s i z e and shape o f c e l l s a r e m i n i m a l and l e s s c o n s i s t e n t . S t u d - i e s r e p o r t e d so f a r i n d i c a t e t h a t s p e c i f i c g r a v i t y i s t h e c h a r a c t e r i s t i c most s e n s i t i v e t o f e r t i l i z i n g and t h a t a maximum o f 10 p e r c e n t r e d u c t i o n c a n o c c u r , b u t t h a t i t i s u s u a l l y more t h a n c o m p e n s a t e d f o r by t h e i n c r e a s e i n volume p r o d u c t i o n (44, 71, 1 1 7 ) . Any d e c r e a s e i n s p e c i f i c g r a v i t y o f t h e raw m a t e r i a l p r o d u c e d by f e r t i l i z a t i o n w i l l be o f g r e a t e s t i m p o r t a n c e i f t h a t m a t e r i a l i s t o be u t i l i z e d by t h e p u l p and p a p e r i n d u s t r y s i n c e t h e y i e l d p e r u n i t volume w i l l be r e d u c e d . A d d i t i o n a l l y , and p e r h a p s l e s s a p p r e c i a t e d , i s t h e e f f e c t a change i n d e n s i t y w o u l d have on t h e p r o p e r t i e s o f t h e p r o d u c t . T h i s c o u l d be b r o u g h t a b o u t b o t h by r e s u l t i n g c hanges i n e i t h e r t h e p r o p o r t i o n 30. o f l a t e w o o d f i b r e s i n t h e raw m a t e r i a l , o r t h r o u g h c h a n g e s i n c e l l m o r p h o l o g y , and p o s s i b l y c e l l w e i g h t . D e p e n d i n g u p on t h e t y p e o f p r o d u c t b e i n g p r o d u c e d t h e c h a n g e s c o u l d e i t h e r be a d v a n t a g e o u s o f d e l e t e r i o u s . The i n f l u e n c e o f s u c h c h a n g e s on t h e q u a l i t y o f t h i s m a t e r i a l when u t i l i z e d as l u m b e r i s o f l e s s i m p o r t a n c e , n o t b e c a u s e t h e s t r e n g t h o f t h e p r o d u c t c o u l d be a l t e r e d b u t , w i t h c u r r e n t g r a d i n g p r a c t i c e s , t h e s e f a c t o r s w o u l d n o t be l i k e l y t o a f f e c t l u m b e r g r a d e . I f and when m a c h i n e g r a d i n g o f s t r u c t u r a l l u m b e r becomes i m p o r t a n t , c h a n g e s i n wood q u a l i t y b r o u g h t a b o u t by f e r t i l i z a t i o n c o u l d p o s s i b l y h a v e a s i g n i f i c a n t e f f e c t on l u m b e r g r a d e ( 6 6 ) . N o t a l l f e r t i l i z e r e x p e r i m e n t s , h o w e v e r , h a v e b e e n s u c c e s s f u l i n o b t a i n i n g a s u b s t a n t i a l and c o n s i s t e n t g r o w t h r e s p o n s e ( 9 2 , 9 3 ) . C o n s e q u e n t l y , i t i s i m p e r a t i v e t o l e a r n more a b o u t t h e b a s i c f a c t o r s a f f e c t i n g t h e q u a n t i t y a n d q u a l i t y o f wood g r o w n i n f e r t i l i z e d s t a n d s . F o r e x a m p l e , .^Zobel et. a l . (173) f o u n d t h a t c e r t a i n l e v e l s o f NPK f e r t i l i z e r t r e a t m e n t h a d d r a s t i c e f f e c t s on s p e c i f i c g r a v i t y o f l o b l o l l y p i n e wood. The same a u t h o r s , as w e l l as P o s e y ( 1 0 8 ) , showed t h e e x i s t e n c e o f d i f f e r e n t i a l e f f e c t s o f f e r t i l i z e r t r e a t m e n t d e p e n d i n g o n i n i t i a l p r o p e r t i e s o f wood. P o s e y a l s o f o u n d t h a t c e r t a i n f o r m u l a t i o n s o f f e r t i l i z e r b r o u g h t g r e a t e r d i f f e r e n c e i n wood p r o p e r t i e s and g r o w t h r a t e c o m p a r e d t o o t h e r s . E a r l i e r e x p l o r a - t o r y w o r k by S a s t r y (117) i n d i c a t e d t h e p o s s i b i l i t y o f d i f f e r e n - t i a l e f f e c t s o f f e r t i l i z e r c o m p o s i t i o n on p h y s i c a l and m e c h a n i - c a l p r o p e r t i e s ' o f D o u g l a s - f i r wood. The p o s s i b i l i t y , e x i s t s t h a t , i n s t u d i e s o f f e r t i l i z e r i n f l u e n c e on o l d e r t r e e s , i . e . , i n f i e l d t r i a l s , s e v e r a l f a c - t o r s r e l a t i n g t o e n v i r o n m e n t and m i c r o - s i t e may c o n f o u n d t h e r e a l e f f e c t s o f f e r t i l i z e r a p p l i c a t i o n ; t h i s i s p a r t o f t h e r e a s o n why some o f t h e t r i a l s may have f a i l e d t o e l i c i t an i n c r e a s e d g r o w t h r e s p o n s e . Such a c o n f o u n d i n g i n f l u e n c e c o u l d p o s s i b l y be c o n t r o l l e d i n l a b o r a t o r y e x p e r i m e n t s e m p l o y i n g s e e d l i n g s . Thus Murphey e t a l . (96) c o n d u c t e d e x p e r i m e n t s i n c o n t r o l l e d n u t r i e n t c u l t u r e s , f o r two g r o w i n g s e a s o n s , u s i n g w e s t e r n l a r c h ( L a r i x o c c i d e n t a l i s ( N u t t . ) ) s e e d l i n g s . T h e i r r e s u l t s i n d i c a t e d i f f e r e n c e s i n s e l e c t e d a n a t o m i c a l f e a t u r e s s u c h as c e l l s i z e and d i m e n s i o n s , number o f t r a c h e i d s p e r 2 10,000 ym , r e s i n c a n a l d i a m e t e r and number o f e p i t h e l i a l c e l l s p e r r e s i n c a n a l , as w e l l as p h y s i c a l and m e c h a n i c a l p r o p e r t i e s . Changes i n c h e m i c a l c o m p o s i t i o n f o l l o w i n g f e r t i l i z a - t i o n have b e e n s t u d i e d t o a v e r y l i m i t e d d e g r e e . L i t t l e s i g n i - f i c a n t change was r e p o r t e d i n D o u g l a s - f i r by E r i c k s o n and L a m b e r t ( 4 4 ) , b u t a s l i g h t l y l o w e r h o l o c e l l u l o s e and a l p h a - c e l l u l o s e c o n t e n t was f o u n d i n t h e wood f o r m e d a f t e r f e r t i l i - z a t i o n . L i k e w i s e , i n y o u n g l o b l o l l y p i n e , c e l l u l o s e y i e l d showed a d e c r e a s i n g t r e n d , b u t t h e change was n o t s i g n i f i c a n t ( 1 7 3 ) . T h e s e e x p e r i m e n t a l r e s u l t s w o u l d be e x p e c t e d . F e r t i l i - z a t i o n has b e e n shown t o i n c r e a s e t h e p r o d u c t i o n o f e a r l y w o o d more t h a n t h a t o f l a t e w o o d , so t h a t t h e g r e a t e r number o f e a r l y w o o d c e l l s w i t h t h e i r h i g h e r l i g n i n c o n t e n t w o u l d d e c r e a s e c e l l u l o s e p e r c e n t a g e and i n c r e a s e t h a t o f l i g n i n . T h e s e c o n - c l u s i o n s a r e i n g e n e r a l a g r e e m e n t w i t h E r i c k s o n ' s (42) f i n d i n g s on i n c r e a s e i n t h e l i g n i n c o n t e n t o f f e r t i l i z e d s p r u c e ( P i c e a a b i e s (L.) K a r s t ) , and Malm's (88) d a t a on t h e i n c r e a s e i n m i d d l e l a m e l l a t h i c k n e s s o f l a t e w o o d c e l l s i n f e r t i l i z e d t r e e s . J e n s e n e t a l . (64) have s t u d i e d t h e e f f e c t o f i n c - r e a s e d g r o w t h r a t e o f p i n e and s p r u c e i n F i n l a n d on wood q u a l i t y , p a r t i c u l a r l y f o r p u l p and p a p e r . I t was r e p o r t e d t h a t p a p e r p r o p e r t i e s were l i t t l e a f f e c t e d by t h e i n c r e a s e d g r o w t h r a t e due t o f e r t i l i z a t i o n . S t u d i e s i n Sweden (51) have r e p o r t e d t h a t p a p e r p r o d u c e d f r o m wood a f t e r f e r t i l i z a t i o n was h i g h e r i n b u r s t and t e n s i l e s t r e n g t h and l o w e r i n t e a r t h a n t h e c o r r e s p o n d i n g m a t e r i a l f r o m u n f e r t i l i z e d t r e e s . Wood o b t a i n e d a f t e r f e r t i l i z a t i o n a l s o p r o d u c e d a d e n s e r and l e s s b u l k y p a p e r . . Changes i n p a p e r c h a r a c t e r i s t i c s c a n be e x p l a i n e d r e a d i l y on t h e b a s i s o f t h e known changes i n wood characteris^„- t i c s due t o f e r t i l i z a t i o n . T hus, t h i n - w a l l e d f i b r e s o f l a r g e d i a m e t e r , s u c h as f o u n d i n f e r t i l i z e d wood i n h i g h p r o p o r t i o n , c o l l a p s e t o f l a t t e n e d r i b b o n s i n t h e c o n v e r s i o n p r o c e s s , t o f o r m a compact, d e n s e s h e e t , low i n o p a c i t y , u n i f o r m i n f o r - m a t i o n , and smooth o f s u r f a c e . F u r t h e r m o r e , t h e y p r e s e n t h i g h l y e f f e c t i v e f i b r e t o f i b r e c o n t a c t a r e a s , r e s u l t i n g i n s t r o n g b o nds, as r e v e a l e d i n h i g h t e n s i l e and b u r s t i n g s t r e n g t h s ( 1 3 ) . S i m i l a r l y , f i b r e l e n g t h i s a d o m i n a n t f a c t o r c o n t r o l l i n g t h e t e a r i n g r e s i s t a n c e o f p a p e r , l o n g f i b r e s p r o d u c i n g p a p e r w i t h t h e h i g h e s t t e a r i n g r e s i s t a n c e by d i s t r i b u t i n g o v e r a l a r g e a r e a t h e s h e a r i n g f o r c e s i n v o l v e d i n t e a r i n g p a p e r ( 1 3 ) . B o t h b u l k and p o r o s i t y i n c r e a s e t o some e x t e n t w i t h i n c r e a s i n g t r a c h e i d l e n g t h ( 2 9 ) . I n summary, i s c a n be s t a t e d t h a t t h e r e s p o n s e o f c o n i f e r o u s t r e e s t o f e r t i l i z a t i o n a p p e a r s t o be r e l a t e d t o t h e v i g o r o f t h e t r e e , t h e i n i t i a l l e v e l o f t h e p a r t i c u l a r wood p r o p e r t y , t h e g e n e t i c a l l y c o n t r o l l e d a b i l i t y o f . t h e i n d i - v i d u a l t o r e s p o n d a n d , f i n a l l y , t h e i n t e r p l a y o f e n v i r o n m e n t . E x t r e m e l y s l o w g r o w i n g t r e e s w h i c h a r e p r o d u c i n g l o w d e n s i t y ' s t a r v a t i o n . w o o d 1 may i n c r e a s e i n d e n s i t y ( 5 1 ) , w h i l e t r e e s g r o w i n g a t a m o d e r a t e r a t e a r e g e n e r a l l y f o u n d t o d e c r e a s e i n d e n s i t y . The m o s t v i g o r o u s t r e e s g r o w i n g c l o s e t o t h e i r p o t e n - t i a l o f t e n show no r e s p o n s e t o f e r t i l i z a t i o n . The d e n s i t y c h a n g e s o b s e r v e d s o f a r a r e f o r t h e m o s t p a r t (up t o 50 p e r c e n t ) a c c o u n t e d f o r by c h a n g e s i n l a t e w o o d p e r c e n t a g e ; b u t some s t u d i e s h a v e i n d i c a t e d t h a t c h a n g i n g r a d i a l w a l l t h i c k n e s s ( m o s t l y l a t e w o o d ) may p l a y some p a r t i n t h i s d e n s i t y r e s p o n s e . I n no s t u d y , h o w e v e r , w e r e c h a n g e s i n s p e c i f i c g r a v i t y r e l a t e d t o c h a n g e s i n c h e m i c a l p r o p e r t i e s o f f e r t i l i z e d wood. L i k e w i s e , a d e c r e a s e i n c e l l l e n g t h i s e x p e c t e d s i n c e t h i s i s a s s o c i a t e d w i t h t h e v e r y r a p i d i n c r e a s e i n g r o w t h , w h i c h c o u l d c a u s e an i n c r e a s e i n t h e number o f c a m b i a l i n i t i a l s due t o c h a n g e i n t h e i r p a t t e r n o f d i v i s i o n . B a n n a n (8) showed t h a t a n t i c l i n a l d i v i s i o n s a r e t h e means by w h i c h m u l t i p l i c a t i o n o f f u s i f o r m c a m b i a l c e l l s i s a c h i e v e d . F r e q u e n c y o f m u l t i - p l i c a t i v e d i v i s i o n s w o u l d i n f l u e n c e t h e l e n g t h o f d e r i v e d i n i t i a l s i n t h e cambium, h e n c e , l e n g t h o f d e r i v e d v a s c u l a r e l e m e n t s ; a f a c t o r s u c h as f e r t i l i z a t i o n , w h i c h w o u l d c a u s e an i n c r e a s e i n r a d i a l g r o w t h , s h o u l d c a u s e a d r o p i n t h e mean c e l l l e n g t h . T h i s c o n c e p t i s s u p p o r t e d by t h e work o f s e v e r a l a u t h o r s (71, 108, 117, 141, 1 7 3 ) . F i n a l l y , as s t a t e d by Z o b e l e t al_. (173) , t h e r e i s i n d i c a t i o n o f a d i f f e r e n t i a l e f f e c t o f f e r t i l i z e r d e p e n d i n g on t h e i n i t i a l wood p r o p e r t y o f t h e t r e e . A l s o , t h e r e a p p e a r s t o be a l a r g e amount o f i n d i v i d u a l t r e e d i f f e r e n c e i n r e s p o n s e t o f e r t i l i z e r t r e a t m e n t s , s u g g e s t i n g g e n e t i c i n t e r a c t i o n . T h i s ' i n d i v i d u a l r e s p o n s e ' may become v e r y i m p o r t a n t i n t h e d e v e l o p - ment o f i m p r o v e d s t r a i n s o f t r e e s . CHAPTER I I I MATERIALS AND METHODS S e l e c t i o n o f M a t e r i a l The p r i m a r y o b j e c t i v e was t o i n c l u d e as w i d e a r a n g e o f t r a c h e i d l e n g t h s as p o s s i b l e , so t h a t t h e c r i t e r i o n f o r t h e s e l e c t i o n o f wood m a t e r i a l was t r a c h e i d l e n g t h b a s e d on measure- ment o f l e n g t h f r o m m a c e r a t e d m a t e r i a l . W i t h i n t h i s o b j e c t i v e , a b r o a d t a x o n o m i c a l s o r t i n g o f wood t i s s u e was made, s o t h a t c o n c l u s i o n s f r o m t h e work m i g h t be i n t e r p r e t e d o V e r a r e a s o n a b l e r a n g e and more c l o s e l y a p p r o a c h a g e n e r a l s t a t e m e n t . I n o r d e r t o i n c l u d e t h e w i t h i n - t r e e v a r i a t i o n i n t r a c h e i d l e n g t h f r o m p i t h t o b a r k , s e v e r a l p o i n t s were s a m p l e d i n one d i s k , a t stump h e i g h t , o f a 5 0 0 - y e a r - o l d D o u g l a s - f i r ( P s e u d o t s u g a men- z i e s i i ( M i r b . ) F r a n c o ) w h i c h was a v a i l a b l e f r o m a n o t h e r s t u d y . The r a d i a l d e n s i t y p a t t e r n i n t h i s d i s k has b een d e s c r i b e d by Kennedy and W a r r e n ( 7 0 ) . From P i n a c e a e , t h r e e s p e c i e s f r o m s e p a r a t e g e n e r a were s a m p l e d : s u g a r p i n e ( P i n u s l a m b e r t i a n a D o u g l . ) , S i t k a s p r u c e ( P i c e a s i t c h e n s i s (Bong.) C a r r . ) and D o u g l a s - f i r . One s p e c i e s o f a r e p r e s e n t a t i v e genuss f r o m e a c h o f t h e s i x o t h e r c o n i f e r o u s f a m i l i e s were i n c l u d e d ; t h e s e were redwood ( S e q u o i a s e m p e r v i r e n s (D. Don) E n d l . ) f r o m T a x o d i a c e a e , New Z e a l a n d w h i t e p i n e ( P o d o c a r p u s d a c r y d i o d e s A. R i c h ) f r o m P o d o c a r p a c e a e , e a s t e r n r e d c e d a r ( J u n i p e r u s v i r g i n i a n a L.) f r o m C u p r e s s a c e a e , P a c i f i c yew (Taxus b r e v i f o l i a N u t t . ) f r o m T a x a c e a e , T a i w a n c o w ' s - t a i l p i n e ( C e p h a l o t a x u s w i l s o n i a n a Hay.) f r o m C e p h a l o - t a x a c e a e , and hoop p i n e ( A r a u c a r i a c u n n i n g h a m i i Sweet) f r o m A r a u c a r i a c e a e . I n a d d i t i o n , two i n c r e m e n t s s h o w i n g p r o n o u n c e d c o m p r e s s i o n wood were a l s o i n c l u d e d f r o m a t r e e o f D o u g l a s - f i r . M a t e r i a l i n t h e f o r m o f i n c r e m e n t c o r e s f r o m f e r - . t i l i z e d D o u g l a s - f i r t r e e s was a v a i l a b l e f r o m a n o t h e r s t u d y ( 1 1 9 ) . Two i n c r e m e n t s , f r o m one t r e e e a c h o f t h r e e f e r t i l i z e r t r e a t m e n t s , were s e l e c t e d f r o m t h i s s t u d y m a t e r i a l . The t r e a t - ments c o n s i s t e d o f f o r m u l a t i o n s o f U r e a , NPK, and ( N H ^ ) ^ w h i c h g e n e r a t e d , r e s p e c t i v e l y , an i n c r e a s e (Urea) and d e c r e a s e (NPK, (NH^^SO^) i n wood s p e c i f i c g r a v i t y f o l l o w i n g a p p l i c a t i o n . F i n a l l y , t r a c h e i d w e i g h t v a r i a t i o n b e f o r e and a f t e r f e r t i l i - z a t i o n was t r a c e d i n s i x c o n s e c u t i v e i n c r e m e n t s o f a D o u g l a s - f i r f e r t i l i z e d w i t h NH^NO^. I n t h i s t r e e , s a m p l i n g i n c l u d e d one g r o w t h i n c r e m e n t b e f o r e f e r t i l i z a t i o n and f i v e i n c r e m e n t s f o r m e d c o i n c i d e n t w i t h and a f t e r t h e two a p p l i c a t i o n s . The e n t i r e s a m p l i n g p l a n i n c l u d e d n i n e g e n e r a , and n i n e s p e c i e s , and a l l s e v e n c o n i f e r o u s f a m i l i e s . A t o t a l o f 28 g r o w t h i n c r e m e n t s were s a m p l e d . Wood m a t e r i a l o f s u g a r p i n e , redwood, hoop p i n e , New Z e a l a n d w h i t e p i n e , and e a s t e r n r e d c e d a r was a v a i l a b l e f r o m t h e x y l a r i u m o f t h e F a c u l t y o f F o r e s t r y , U n i v e r s i t y o f B r i t i s h C o l u m b i a . A s m a l l wood sample o f C e p h a l o - t a x u s , i n t h e f o r m o f a d i s k , was o b t a i n e d f r o m N a t i o n a l T a i w a n U n i v e r s i t y , E x p e r i m e n t a l R e s e a r c h F o r e s t , i n T a i w a n . The . - r e m a i n i n g m a t e r i a l was c o l l e c t e d w i t h i n B r i t i s h C o l u m b i a by t h e a u t h o r , f r o m t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a R e s e a r c h F o r e s t , n e a r Haney and R o b e r t s o n R i v e r V a l l e y , S o u t h C e n t r a l V a n c o u v e r I s l a n d . O r i g i n , d e s c r i p t i o n o f g r o s s wood c h a r a c - t e r i s t i c s , and measured t r a c h e i d l e n g t h o f t h e s e wood sam p l e s a r e g i v e n i n A p p e n d i x 1. Wood Sample and T r a c h e i d S k e l e t o n P r e p a r a t i o n G e n e r a l p r o c e d u r e : S e l e c t e d wood g r o w t h i n c r e m e n t s were e x c i s e d f r o m wood b l o c k s , d i s k s o r i n c r e m e n t c o r e s , t h e n c a r e f u l l y s e p a r a t e d i n t o w h o l e e a r l y w o o d and l a t e w o o d p o r t i o n s ; e a c h o f t h e s e was t h e n d i s s e c t e d i n t o s m a l l e r p i e c e s , w i t h p o r t i o n s b u n d l e d s e p a r a t e l y i n s t a i n l e s s s t e e l w i r e s c r e e n (150-mesh) and made i n t o s m a l l p a c k e t s ; e a c h p a c k e t was l a b e l l e d w i t h a m e t a l t a g t o i d e n t i f y t h e s p e c i e s , i n c r e m e n t number, and e a r l y w o o d and l a t e w o o d f r a c t i o n s . S e p a r a t e p a c k e t s o f some o f t h e i n c r e m e n t s were a l s o p r e p a r e d t o d e t e r m i n e t h e p e r c e n t a g e y i e l d o f h o l o c e l l u l o s e and a l p h a - c e l l u l o s e . H o l o c e l l u l o s e p r e p a r a t i o n : P a c k e t s c o n t a i n i n g wood p i e c e s were e x t r a c t e d w i t h 2:1 (V/V) e t h a n o l - b e n z e n e ; h o l o - c e l l u l o s e t r a c h e i d s k e l e t o n s were o b t a i n e d by t r e a t i n g wood p i e c e s ( i n p a c k e t s ) i n p e r a c e t i c a c i d / w i t h s u b s e q u e n t s o d i u m b o r o h y d r i d e r e d u c t i o n , f o l l o w i n g t h e method d e s c r i b e d by L e o p o l d ( 8 3 ) . T h i s method i s r e p o r t e d t o g i v e v i r t u a l l y i n t a c t s u p e r i o r h o l o c e l l u l o s e f i b r e s w i t h 93 t o 100 p e r c e n t r e c o v e r y o f m a j o r 38. p o l y s a c c h a r i d e s and minimum c a r b o x y l g r o u p s . The p r o c e d u r e i n c l u d e d t h r e e s t e p s : 1. The t r e a t m e n t was d i v i d e d i n t o a number o f 3 0 m i n u t e c y c l e s , c o n s i s t i n g o f a p e r a c e t i c a c i d t r e a t m e n t ( i . e . , 5 g o f p e r a c e t i c a c i d and 2.5 g o f s o d i u m a c e t a t e p e r gram o f wood; t o t a l v olume 32 ml p e r gram o f wood; t e m p e r a t u r e 7 0 ° C ) , f o l l o w e d by s o a k i n g i n h o t w a t e r (30 m i n u t e s , t e m p e r a t u r e 7 0 ° C ) . 2. A t t h e end o f e a c h c y c l e (3 o r 5; s e e below) t h e p a c k e t s were washed i n d i s t i l l e d w a t e r and t h e w a t e r removed w i t h a c e t o n e . A f t e r a i r - d r y i n g , t h e p a c k e t s were t h e n p l a c e d i n a s u c t i o n f l a s k , t h e f l a s k was e v a c u a t e d , and a new b a t c h o f r e - a g e n t was drawn i n t o i t . 3. A t t h e c o n c l u s i o n o f t h e s o d i u m b o r o h y d r i d e r e d u c - t i o n (3 h r , pH 9.5), t h e d e l i g n i f i e d , s o f t , wood p i e c e s were d e f i b e r i z e d by g e n t l y r u b b i n g w i t h a g l a s s r o d a g a i n s t t h e w a l l o f a g l a s s b e a k e r i n t h e p r e s e n c e o f h o t w a t e r . A t o t a l o f f i v e c y c l e s was needed f o r l a t e w o o d and t h r e e c y c l e s f o r e a r l y w o o d d e l i g n i f i c a t i o n as d e t e r m i n e d by p h y s i c a l c h e c k s . The h o l o c e l l u l o s e t r a c h e i d s k e l e t o n s t h u s o b t a i n e d were s t o r e d i n t e s t t u b e s c o n t a i n i n g d i s t i l l e d w a t e r . • A l p h a - c e l l u l o s e p r e p a r a t i o n : A p p r o x i m a t e l y h a l f t h e q u a n t i t y o f t h e h o l o c e l l u l o s e f r a c t i o n , f r o m e a c h s a m p l e , was f u r t h e r r e d u c e d t o a l p h a - c e l l u l o s e s k e l e t o n s e s s e n t i a l l y f o l l o w i n g T A P P I s t a n d a r d m e thod T203 o s - 6 1 (131) s c a l e d t o accommodate m i c r o - a m o u n t s o f m a t e r i a l . C o n v e r t e d y i e l d s o b t a i n e d f o r h o l o - a n d a l p h a - c e l l u l o s e f r a c t i o n s a r e g i v e n s e p a r a t e l y f o r e a r l y w o o d and l a t e w o o d i n A p p e n d i x I I . T r a c h e i d s k e l e t o n s o f a l p h a - c e l l u l o s e , a f t e r w a s h i n g , w e r e a l s o k e p t s t o r e d i n t e s t t u b e s c o n t a i n i n g d i s t i l l e d w a t e r . A t a l l s t a g e s h o l o - and a l p h a - c e l l u l o s e p u l p s w e r e k e p t i n w e t c o n d i - t i o n a n d n e v e r a l l o w e d t o d r y . T h i s was done s o t h a t t r a c h e i d l e n g t h m e a s u r e m e n t s c o u l d be b a s e d on g r e e n c o n d i t i o n and r e p r e s e n t more c l o s e l y t h e i r n a t u r a l s t a t e i n t h e t r e e . T h u s , any p o s s i b l e e f f e c t s o f s h r i n k a g e due t o d r y i n g w e r e a v o i d e d as f a r as p o s s i b l e . The o n l y e x c e p t i o n s w e r e t h o s e s a m p l e s w h i c h w e r e i n t e n d e d f o r y i e l d d e t e r m i n a t i o n . T h e s e p u l p s ( h o l o c e l l u l o s e ) w e r e p r o c e s s e d i n t o t h i n s h e e t s a nd a i r - d r i e d , b e f o r e a l k a l i t r e a t m e n t . T e s t f o r l i g n i n : L i g n i n c o n t e n t i n some o f t h e h o l o - c e l l u l o s e and a l p h a - c e l l u l o s e s a m p l e s was d e t e r m i n e d by T A P P I s t a n d a r d m e thod T236 m-60 (130) as m o d i f i e d f o r r e d u c e d q u a n - t i t y o f p u l p by B e r z i n s ( 1 2 ) . The t e s t showed t h a t r e s i d u a l l i g n i n c o n t e n t s w e r e e q u a l t o o r l e s s t h a n 1.53 and 0.29 p e r c e n t , r e s p e c t i v e l y , f o r h o l o - a n d a l p h a - c e l l u l o s e , i n t h e s a m p l e s e x a m i n e d ( s e e A p p e n d i x I I ) . 40. A n a l y s i s o f T r a c h e i d S k e l e t o n s 1. S a m p l i n g c r i t e r i o n The p r i m a r y o b j e c t i v e o f t h i s i n v e s t i g a t i o n was t o r e l a t e t r a c h e i d w e i g h t t o i t s l e n g t h . I n i t i a l l y , a l l t r a c h e i d s were t r e a t e d as b e l o n g i n g t o one p o p u l a t i o n , i r r e s p e c t i v e o f s o u r c e . I n o r d e r t o examine s u c h a r e l a t i o n s h i p t h e sample r e p l i c a t i o n number n e e d e d t o be d e t e r m i n e d . C a l c u l a t i o n o f sample number f o r a s t a t i s t i c a l l y d e f e n s i b l e d e t e r m i n a t i o n c o u l d be o b t a i n e d i f t h e o b j e c t i v e were t o e s t a b l i s h 'norms' f o r i n c r e m e n t s o r t i s s u e f r a c t i o n w i t h i n i n c r e m e n t s . However, i n a s t u d y s u c h as t h i s where s a m p l i n g i n c l u d e d a w i d e r a n g e o f t r a c h e i d l e n g t h s , f r o m d i f f e r e n t o r i g i n s , t h e p r i m a r y f u n c t i o n i s n o t t o e s t i m a t e i n c r e m e n t mean; r a t h e r i t i s t o t e s t t h e o v e r a l l r e l a t i o n s h i p b etween t h e two c h a r a c t e r i s t i c s , namely t r a c h e i d l e n g t h and w e i g h t . M o r e o v e r , w i t h i n - t r e e t r a c h e i d l e n g t h p a t t e r n s , and t h o s e between and w i t h i n s p e c i e s o f c o n i - f e r s , h a v e a l r e a d y b e e n e s t a b l i s h e d by s e v e r a l w o r k e r s . T h e r e - f o r e , s t a n d a r d p r o c e d u r e s s u c h as c a l c u l a t i o n o f sample s i z e by S t e i n ' s t w o - s t a g e sample p r o c e d u r e ( 1 2 5 ) , a r e o f l i t t l e h e l p . S t i l l , i n o r d e r t o m a x i m i z e r e s e a r c h e f f o r t w i t h i n t h e t i m e a v a i l a b l e , i t i s e s s e n t i a l t o d e t e r m i n e sample s i z e s o as t o g e t r e l i a b l e e s t i m a t e s o f t h e d e s i r e d r e l a t i o n s h i p . P r e v i o u s e x p e r i e n c e f r o m a s i m i l a r s t u d y (118) i n d i c a t e d t h a t a t l e a s t two g r o u p s o f 10 f i b r e s e a c h , t a k e n a t random f r o m t h e e a r l y - wood and l a t e w o o d r e g i o n s o f t h e i n c r e m e n t , w o u l d p r o v i d e a s a t i s f a c t o r y number w i t h r e a s o n a b l e e s t i m a t e s o f t h e mean and s t a n d a r d d e v i a t i o n s . F o r t h e above r e a s o n s , t h e number o f t r a c h e i d s a n a l y z e d f r o m one p a r t i c u l a r sample ( i n c r e m e n t ) was s t a n d a r - d i z e d a t a p p r o x i m a t e l y 20. I n some c a s e s where t h e v a r i a t i o n i n t r a c h e i d l e n g t h was g r e a t , t h i s number was i n c r e a s e d e v e n up t o 40 f o r t h a t s a m p l e . 2. T r a c h e i d l e n g t h A p o r t i o n o f t h e m a t e r i a l f r o m e a c h o f t h e l a b e l l e d t e s t t u b e s was t r a n s f e r r e d t o a p e t r i d i s h ; r a n d o m l y s e l e c t e d t r a c h e i d s were p i c k e d , w i t h t h e h e l p o f f i n e t w e e z e r s , u s i n g a b i n o c u l a r m i c r o s c o p e . T h e s e were t h e n t r a n s f e r r e d t o a g l a s s s l i d e i n a d r o p o f d i s t i l l e d w a t e r and a c o v e r g l a s s a p p l i e d . C a r e was t a k e n t h a t no damage was done t o t h e f i b r e t i p s . T r a c h e i d l e n g t h measurements were made, i n t h e wet c o n d i t i o n , by t h e u s e o f a f i b r e l e n g t h r e c o r d e r . Images o f f i b r e s a r e p r o j e c t e d on an opaque g l a s s s c r e e n and t h e i r l e n g t h s m e a s u r e d w i t h t h e a i d o f a c a l i b r a t e d p r o b e w h e e l . T h i s i n s t r u m e n t has- va p r e c i s i o n o f + 0.1042 mm a t 56.X and + 0.0712 mm a t 82 X m a g n i f i c a t i o n . T r a c h e i d s up t o 4.9 8 mm l e n g t h were m e a s u r e d u s i n g t h e h i g h e r m a g n i f i c a t i o n . As n o t e d , a minimum o f 10 t r a c h e i d s wass. m e a s u r e d s e p a r a t e l y f o r e a r l y w o o d and l a t e w o o d f o r e a c h o f t h e s e l e c t e d i n c r e m e n t s f r o m e a c h o f t h e h o l o - andv a l p h a - c e l l u l o s e p u l p s . 42. A l p h a - c e l l u l o s e t r a c h e i d s k e l e t o n s , showed a s h o r t e n i n g i n l e n g t h o f up t o 15 p e r c e n t i n r e l a t i o n t o h o l o c e l l u l o s e t r a c h e i d s k e l e t o n s . T h i s was f u r t h e r v e r i f i e d by t r e a t i n g i n d i v i d u a l t r a c h e i d s k e l e t o n s o f h o l o c e l l u l o s e w i t h a 17.5 p e r c e n t NaOH s o l u t i o n i n s e p a r a t e p e t r i d i s h e s , f o r 75 m i n u t e s a t 20°C. L e n g t h measurements b e f o r e and a f t e r t h e t r e a t m e n t c o n f i r m e d t h e r e d u c t i o n i n t r a c h e i d l e n g t h . The a l k a l i n e t r e a t m e n t o f h o l o c e l l u l o s e f i b r o u s s k e l e t o n s c o u l d be e x p e c t e d t o c a u s e s w e l l i n g (21) due t o changes i n l a t t i c e and t h e r e b y s h o r t e n i n g o f t h e t r a c h e i d , e x p l a i n a b l e by s i m p l e m e c h a n i c a l f a c t o r s , i . e . , r e v e r s e P o i s s o n ' s r a t i o . . ( 1 6 0 ) . 3. T r a c h e i d w e i g h t A f t e r d e t e r m i n i n g t h e l e n g t h , e a c h t r a c h e i d was t r a n s f e r r e d t o a s e p a r a t e l a b e l l e d p l a s t i c w e i g h b o a t . S e v e r a l o f t h e s e b o a t s were t h e n p l a c e d i n a f r e e z e d r y e r ( V i r t i s M o d e l 10-100) and d r i e d i n vacuum (-60°C and 5 u vacuum) f o r f o u r h o u r s . L a t e r e x p e r i e n c e showed t h a t one h o u r was s u f f i c i e n t t o d r y i n d i v i d u a l f i b r e s . S u b s e q u e n t l y t h e b o a t s , e a c h w i t h i t s f i b r e , were t r a n s f e r r e d t o a c o n s t a n t t e m p e r a t u r e and h u m i d i t y (CTH) room (73 + 3 . 5 ° F , 50 + 2% RH) and c o n d i t i o n e d f o r 24 h o u r s . T r a c h e i d s f o r w h i c h l e n g t h s had been d e t e r m i n e d were t h e n w e i g h e d i n d i v i d u a l l y , u s i n g a s p e c i a l l y d e s i g n e d q u a r t z u l t r a - m i c r o b a l a n c e . B r i e f l y , t h e b a l a n c e c o n s i s t s o f a t h i n q u a r t z f i l a m e n t (20 y d i a m e t e r and 6.35 cm (2 1/2 i n . ) l o n g ) f i r m l y f i x e d a t one e n d . The t r a c h e i d t o be w e i g h e d i s p l a c e d on t h e f r e e end and i t s w e i g h t d e t e r m i n e d by m e a s u r i n g t h e d e f l e c t i o n o f t h e f i l a m e n t w i t h a m i c r o s c o p e . The p r i n c i p l e i s s i m i l a r t o t h e l o a d i n g o f a c a n t i l e v e r beam a t t h e f r e e e n d . The b a l a n c e so d e v e l o p e d has a w e i g h t i n g r a n g e f r o m 0.06 t o 14 y g , 1 and p r o v i d e s a r e p r o d u c i b i l i t y ( p r e c i s i o n ) w i t h i n 0.03 y g , w h i c h i s a b o u t 10 t i m e s h i g h e r t h a n t h a t o f p r e s e n t c o m m e r c i a l e l e c t r o b a l a n c e s and u l t r a - m i c r o b a l a n c e s . The b a l a n c e c a s e c o n s i s t s o f a 20 ml h y p o d e r m i c s y r i n g e , t o p r o t e c t t h e f i l a m e n t , and a mount, i . e . , t h e p l u n g e r , f o r h o l d i n g t h e f i l a m e n t i n p o s i t i o n . The c l o s e d end o f t h e s y r i n g e i s c u t o f f and l e f t open t o make l o a d i n g o f t h e f i l a m e n t p o s s i b l e , w i t h a r r a n g e m e n t s made t o c o v e r t h e o p e n i n g w i t h a c o v e r g l a s s d u r i n g w e i g h i n g s t o p r o t e c t i t f r o m a i r c u r r e n t s . A d i a g r a m o f t h e u l t r a - m i c r o b a l a n c e i s g i v e n b e l o w . A c r u c i a l s t e p i n t h e u s e o f t h e b a l a n c e i s i t s c a l i b r a t i o n . T h i s was a c h i e v e d by a c o l o r i m e t r i c method, i n w h i c h s m a l l c r y s t a l s o r f l a k e s o f f l u o r e s c e i n were l o a d e d on t h e f i l a m e n t and t h e d e f l e c t i o n m e a s u r e d . T h e s e c r y s t a l s were t h e n c o l l e c t e d and d i s s o l v e d i n a known volume o f s o l v e n t (1 p e r c e n t N a ^O^) and t h e o p t i c a l d e n s i t y was m e a s u r e d a g a i n s t a p p r o p r i a t e s t a n d a r d s , u s i n g a Unicam SP 800 R e c o r d i n g S p e c t r o - p h o t o m e t e r a t a b s o r p t i o n maxima o f 493 and 460 m.U R e s u l t s were t h e n compared w i t h t h e o p t i c a l d e n s i t y o f a s e r i e s o f 1 — 6 1 m i c r o gram (yg) = 1 0 g 44. 1 2 6 Cross s e c t i o n of the Ultra-Wicrobalance. 1. g l a s s plunger 2. open-ended g l a s s tube 3. p l a s t i c l o ck t i p 4. p l a s t i c needle mount. 5. quartz f i l a m e n t 6. cover g l a s s l i d 7. radium f o i l s o l u t i o n s t h e s t r e n g t h s o f w h i c h were known w i t h i n c l o s e t o l e r a n c e l i m i t s . The f i n a l . r e s u l t was a g r a p h s h o w i n g de- f l e c t i o n o f t h e f i l a m e n t a g a i n s t w e i g h t o f m a t e r i a l p l a c e d on i t . The r e l a t i o n s h i p between sample w e i g h t and v e r t i c a l d i s - p l a c e m e n t o f t h e f i l a m e n t was l i n e a r o v e r t h e e n t i r e u s e f u l r a n g e o f t h e b a l a n c e . Once c a l i b r a t e d , t h e b a l a n c e . ( u s i n g t h e same q u a r t z f i l a m e n t ) was u s e d t o w e i g h t r a c h e i d s . p e r m i n u t e ; t h e r e s u l t s were r e p r o d u c i b l e when r e w e i g h i n g t h e same i n d i v i d u a l (+ 0.03 u g ) . F u r t h e r d e t a i l s o f t h e b a l a n c e c o n s t r u c t i o n , d e s i g n and c a l i b r a t i o n a r e d i s c u s s e d e l s e w h e r e (118, 1 2 1 ) . W e i g h t s o f some o f t h e t r a c h e i d s (1 t o 2 ug) a r e T r a c h e i d s may be w e i g h e d a t t h e r a t e o f one o r two c o m p a r a b l e (+ 0.15 yg) t o t h o s e o b t a i n e d by u s i n g a M e t t l e r UM7 b a l a n c e , as d e t e r m i n e d by c h e c k s . C o r r e c t i o n f o r m o i s t u r e c o n t e n t : The w e i g h t o f t r a c h e i d s t h u s o b t a i n e d r e p r e s e n t s t h a t a t t h e EMC o f t h e CTH room. I n o r d e r t o e x p r e s s t h e w e i g h t i n o v e n d r y c o n d i t i o n , a d j u s t m e n t s were'made t o t h e o r i g i n a l w e i g h t . T h i s was done as f o l l o w s : t h e e x t r a m a t e r i a l a v a i l a b l e f r o m t h e h o l o - and a l p h a - c e l l u l o s e p u l p s ( a f t e r s e l e c t i o n o f t h e r e q u i r e d number o f t r a c h e i d s f o r w e i g h i n g ) was made i n t o t h i n s h e e t s o f f i b r e , and w a t e r removed by a p p l y i n g s u c t i o n . T h e s e " p u l p s h e e t s " were numbered f o r i d e n t i f i c a t i o n , and k e p t s t o r e d ( i n t h e open) i n t h e CTH room u n t i l a c o n s t a n t w e i g h t was r e a c h e d , f o l l o w i n g w h i c h t h e y were o v e n d r i e d . I n d i v i d u a l t r a c h e i d w e i g h t s were t h e n a d j u s t e d t o m o i s t u r e - f r e e b a s i s , t o t h e n e a r e s t 0.01 p e r c e n t . The m o i s t u r e c o n t e n t s o f t h e p u l p s a m p l e s e xamined a r e g i v e n i n A p p e n d i x I I I . S t a t i s t i c a l A n a l y s e s S e v e r a l a n a l y s e s were c o n d u c t e d i n o r d e r t o t e s t t h e p r o p o s e d h y p o t h e s i s and i t s a p p l i c a b i l i t y t o woods o f d i f f e r e n t o r i g i n . M u l t i p l e r e g r e s s i o n a n a l y s e s were u s e d t o d e t e r m i n e i n t e r r e l a t i o n s h i p s w i t h i n and between s p e c i e s , and w i t h i n i n c r e m e n t s ( e a r l y w o o d and l a t e w o o d ) ; t o a s s e s s t h e measure o f a s s o c i a t i o n between t r a c h e i d l e n g t h and w e i g h t ; f i n a l l y , t o d e v e l o p r e g r e s s i o n e q u a t i o n s o f b e s t f i t . I n a d d i t i o n t o t h e l e a s t - s q u a r e s m o d e l s , c o n d i t i o n e d r e g r e s s i o n m o d e l s ( w i t h z e r o i n t e r c e p t ) were a l s o u s e d . The r e g r e s s i o n t e c h n i q u e r e q u i r e d a s t e p w i s e e l i m i n a t i o n p r o c e d u r e , f r o m an e q u a t i o n o f m u l t i p l e v a r i a b l e s , as d e v e l o p e d by. Kozak and S m i t h ( 7 3 ) . A n a l y s e s o f - c o v a r i a n c e w e r e " e m p l o y e d t o t e s t d i f - f e r e n c e s w i t h i n and between s p e c i e s ; b etween wood f o r m e d a t j u v e n i l e , m a t u r e and o v e r m a t u r e s t a g e s o f t r e e g r o w t h ; between e a r l y w o o d and l a t e w o o d ; b e t w e e n n o r m a l wood and c o m p r e s s i o n wood; and t o c h a r a c t e r i z e t h e b e h a v i o u r o f wood f r o m f e r t i l i z e d t r e e s . C o v a r i a n c e i s a t e c h n i q u e u s e d t o a d j u s t means o f t h e d e p e n d e n t v a r i a b l e 'Y' ( T r a c h e i d W e ight) f o r d i f f e r e n c e s i n s e t s o f v a l u e s o f t h e c o r r e s p o n d i n g i n d e p e n d e n t v a r i a b l e 'X' ( T r a c h e i d L e n g t h ) and t o b r i n g o u t d i f f e r e n c e s , i f any, w i t h i n 2 o r b e t w e e n t h e g r o u p s n o t e d a b o v e . The a n a l y s i s e mployed compares r e g r e s s i o n e q u a t i o n s f o r p a r a l l e l i s m and c o i n c i d e n c e . I t s p u r p o s e i s t o d e t e r m i n e w h e t h e r s e p a r a t e r e g r e s s i o n e q u a t i o n s s h o u l d be u s e d f o r e a c h p o p u l a t i o n u n d e r s t u d y a n d / o r i f some o r a l l o f t h e p o p u l a t i o n s c o u l d be d e s c r i b e d by one r e g r e s s i o n - e q u a t i o n . A s i m p l e method d e v e l o p e d by Dr..A. Kozak, F a c u l t y o f F o r e s t r y , The U n i v e r s i t y o f B r i t i s h C o l u m b i a , and p r e s e n t e d a t t h e 3 r d C o n f e r e n c e o f t h e A d v i s o r y Group o f F o r e s t S t a t i s - t i c i a n s , S e c t i o n 25, I.U.F.R.O., F r a n c e , September 7-11, 1970. CHAPTER IV RESULTS AND DISCUSSION The p r e s e n t s t u d y i s n o t c o n c e r n e d w i t h d e s c r i p t i o n o f t r a c h e i d s i n s p e c i e s o f d i f f e r e n t t a x o n o m i c o r i g i n n o r , - f o r t h a t m a t t e r , w i t h d e v e l o p m e n t o f p r o c e d u r e s t o t e s t d i f f e r e n c e s i n t r a c h e i d w e i g h t s w i t h i n t r e e s . R a t h e r , i t s p u r p o s e and a t t e n t i o n a r e f o c u s s e d on: a) e s t a b l i s h i n g a q u a n t i t a t i v e r e l a t i o n s h i p between t r a c h e i d l e n g t h a n d . t h e amount o f c a r b o h y d r a t e f r a c t i o n ( h o l o - and a l p h a - c e l l u l o s e ) t h a t may e x i s t w i t h i n and b etween c o n i f e r o u s s p e c i e s ; b) e v o l v i n g a b a s i c r e l a t i o n s h i p f o r c o n i f e r o u s woods, f r o m b i o l o g i c a l and c e l l m o r p h o l o g i c a l c o n c e p t s , i n a i d o f p r a c t i c a l c o n s i d e r a t i o n s , f o r example c a r b o h y d r a t e y i e l d b a s e d on t r a c h e i d l e n g t h ; and c) t o t e s t t h e u s e f u l n e s s o f s u c h a b a s i c r e l a t i o n s h i p w i t h r e s p e c t t o o b s e r v e d e f f e c t s o f f e r t i l i z a t i o n on wood c h a r a c t e r i s t i c s i n D o u g l a s - f i r . Such a f u n d a m e n t a l r e l a t i o n s h i p f o r c o n i f e r o u s s p e c i e s c o u l d a l s o p r o v i d e means t o , o r r e - e m p h a s i z e t h e i m p o r t a n c e o f , t r a c - h e i d l e n g t h f o r s e l e c t i o n o f i n d i v i d u a l s w i t h i n s p e c i e s i n t r e e i mprovement work. 48. R e l a t i o n s h i p Between T r a c h e i d Length, and W e i g h t . 1. V a r i a t i o n w i t h i n s p e c i e s A t o t a l o f 1,534 i n d i v i d u a l t r a c h e i d s (787 h o l o - c e l l u l o s e s k e l e t o n s and 747 a l p h a - c e l l u l o s e s k e l e t o n s ) was m e a s u r e d f o r l e n g t h and w e i g h t i n t h e n i n e s p e c i e s s e l e c t e d . Mean v a l u e s f o r e a c h o f t h e i n c r e m e n t s e xamined a r e g i v e n s e p a r a t e l y f o r e a r l y w o o d and l a t e w o o d , i n T a b l e s I and I I . T r a c h e i d w e i g h t and l e n g t h v a l u e s , t a k e n r a d i a l l y a c r o s s t h e d i s k s , f o r two D o u g l a s - f i r t r e e s , a r e c o n t a i n e d i n T a b l e s I I I and IV. O n l y t h e d a t a f r o m 'normal wood' ( e x c l u d i n g compres- s i o n wood and wood f r o m f e r t i l i z e d t r e e s ) were c o n s i d e r e d when e s t a b l i s h i n g a r e l a t i o n s h i p between w e i g h t o f f i b r e s k e l e t o n and i t s l e n g t h . I n a d d i t i o n , t h e r e l a t i o n s h i p s l f : o r b o t h h o l o - and a l p h c e l l u l o s e s k e l e t o n s were examined s e p a r a t e l y . An a d v a n t a g e o f knowing t h e amount o f h o l o c e l l u l o s e i s t h a t i t p r o v i d e s an e s t i m a t e ( w i t h i n r e a s o n a b l e l i m i t s ) o f t h e t o t a l t r a c h e i d c a r - b o h y d r a t e f r a c t i o n , whereas a l p h a - c e l l u l o s e r e l a t e s t o t h e b a s i s k e l e t a l component. I n d i v i d u a l s p e c i e s r e l a t i o n s h i p s o b t a i n e d between t r a c h e i d w e i g h t and l e n g t h a r e shown i n F i g u r e s 1 t o 9, a l o n g w i t h t h e r e g r e s s i o n r e l a t i o n s h i p s o f b e s t f i t . I t i s s e e n t h a t t h e c o r r e l a t i o n s d e t e r m i n e d u s i n g e i t h e r t h e c u r v i l i n e a r o r t h e l i n e a r r e l a t i o n s h i p , by l e a s t - s q u a r e s method, a r e h i g h l y s i g n i f i c a n t , a c c o u n t i n g f o r 66.4 t o 94.8 and 84.8 t o 98.6 p e r 49. c e n t o f v a r i a b i l i t y , i n h o l o c e l l u l o s e and a l p h a - c e l l u l o s e s k e l e t o n w e i g h t s , r e s p e c t i v e l y , by t h e l e n g t h f a c t o r a l o n e . The v a r i a b i l i t y a c c o u n t e d f o r by l e n g t h i s h i g h e r f o r a l p h a - c e l l u l o s e t h a n i t i s f o r h o l o c e l l u l o s e . P o s s i b l y t h i s i s r e l a t e d t o t h e f a c t t h a t a l p h a - c e l l u l o s e , b e i n g more b a s i c i n t h e s k e l e t a l s t r u c t u r e o f c a r b o h y d r a t e framework, s h o u l d be l e s s v a r i a b l e . T h e r e c o u l d be two o t h e r p o s s i b i l i t i e s , ~ t h e s e b e i n g : 1. The s w e l l i n g and s h r i n k a g e changes t h a t o c c u r r e d due t o s t r o n g a l k a l i t r e a t m e n t , may have removed some o f t h e m i n u t e v a r i a t i o n s i n l e n g t h - w e i g h t r e - l a t i o n s h i p s f o r t h e s e f i b r e s ; and 2. The t e c h n i q u e o f a l p h a - c e l l u l o s e i s o l a t i o n ( 1 3 1 ) , may be a b e t t e r c o n t r o l l e d one t h a n t h a t o f h o l o - c e l l u l o s e (83) . The s t r o n g r e l a t i o n s h i p s . o b e t w e e n t r a c h e i d l e n g t h and w e i g h t , as o b s e r v e d f r o m F i g u r e s 1 t o 9 and t h e s t a t i s t i c a l a n a l y s e s g i v e n t h e r e i n , a r e h i g h l y s i g n i f i c a n t , f o r e a c h o f t h e n i n e s p e c i e s e x a m i n e d . T h i s i s o f i m p o r t a n c e s i n c e b o t h t r a c h e i d l e n g t h and g r o s s wood c a r b o h y d r a t e c o n t e n t i n c r e a s e w i t h age, i . e . , f r o m p i t h t o w a r d s b a r k , f o r a number o f y e a r s as d e s - " c r i b e d p r e v i o u s l y . R e s u l t s o f g r o s s wood a n a l y s e s done by s e v e r a l w o r k e r s s u g g e s t e d a r e l a t i o n s h i p between c a r b o h y d r a t e c o m p o s i t i o n o f wood and c e l l l e n g t h . B u t , due t o c o n f o u n d i n g i n f l u e n c e o f t h e age e f f e c t , t h e r e l a t i o n s h i p o b t a i n e d was a t t r i b u t e d t o g r o w t h f a c t o r s . D a d s w e l l e t a_l. ( 2 8 ) , who o b t a i n e d a h i g h l y s i g n i f i c a n t c o r r e l a t i o n (1 p e r c e n t l e v e l ) f o r r a d i a t a p i n e , s u g g e s t e d t h a t : t h i s i s due no d o u b t t o t h e i n c r e a s e i n b o t h w i t h age, b u t i t i s a f a c t t h a t i s i m p o r t a n t b e c a u s e f i b e r l e n g t h c a n p o s s i b l y be c o n - t r o l l e d g e n e t i c a l l y , and i t w o u l d be most a d v a n t a g e o u s i f h i g h a v e r a g e f i b e r l e n g t h a l s o means h i g h c e l l u l o s e c o n t e n t . I f j u and Kennedy (60) a l s o o b t a i n e d a h i g h l y s i g n i - f i c a n t ' r ' v a l u e between t r a c h e i d l e n g t h and g r o s s wood c e l l u - l o s e c o n t e n t f o r D o u g l a s - f i r . T h e s e a u t h o r s a l s o e x p l a i n e d t h e c o r r e l a t i o n s o b t a i n e d on t h e b a s i s o f i n c r e a s e w i t h age o f b o t h t r a c h e i d l e n g t h and c e l l u l o s e c o n t e n t i n a s i n g l e stem. I n a d d i t i o n , t h e a u t h o r s o f f e r e d an e x p l a n a t i o n " t h a t more compound m i d d l e l a m e l l a e w i l l o c c u r p e r u n i t o f l e n g t h i n a l o n g i t u d i n a l row o f s h o r t t r a c h e i d s , " w h i c h s h o u l d e x p l a i n t h e o b s e r v e d t r a c h e i d l e n g t h and g r o s s c e l l u l o s e c o n t e n t r e l a t i o n - s h i p . T h i s c o n c e p t was s u p p o r t e d by s t u d i e s o f E i n s p a h r e t a l . (3 8, 39) who n o t e d n e g a t i v e r e l a t i o n s h i p s between p u l p f i b r e l e n g t h and g r o s s wood l i g n i n c o n t e n t , f o r s l a s h p i n e and l o b l o l l y p i n e . They s u g g e s t e d t h a t " l o n g e r f i b r e s c o u l d - r e s u l t i n f e w e r end w a l l s p e r g i v e n w e i g h t o f wood and w o u l d c o n s e q u e n t l y r e s u l t i n l e s s l i g n i n (and l e s s p e c t i n compounds) and a h i g h e r p u l p y i e l d " ; i . e . , i n c r e a s e d f i b r e l e n g t h r e s u l t s i n f e w e r f i b r e s p e r gram and t h i s i n t u r n means l e s s i n t e r c e l l u l a r mat- e r i a l and l e s s l i g n i n . F u r t h e r , t h e y f o u n d t h a t f i b r e l e n g t h and c e l l d i a m e t e r a r e h i g h l y c o r r e l a t e d , f r o m w h i c h t h e y 51.. c o n c l u d e d t h a t t h e r e l a t i v e p r o p o r t i o n o f t h e c e l l w a l l l a y e r s may a l s o change due t o g e n e r a l i n c r e a s e i n c e l l s i z e . I n v i e w o f t h e a g e - r e l a t e d r e l a t i o n s h i p between g r o s s wood c e l l u l o s e c o n t e n t and c e l l l e n g t h w i t h i n a p a r t i c u - l a r stem, t h e q u e s t i o n a r i s e s as t o w h e t h e r a s i m i l a r r e l a t i o n - s h i p h o l d s f o r d i f f e r e n t stems o f one s p e c i e s , and between s p e c i e s , and w h e t h e r i t i s c h a r a c t e r i s t i c o f a s p e c i e s . One a p p r o a c h c a n be f o u n d i n t h e work o f Z o b e l e t a l . (17 2 ) . \ T h e s e a u t h o r s n o t e d a b e t w e e n - t r e e c o r r e l a t i o n f o r t r a c h e i d l e n g t h a t 30 y e a r s o f age and p e r c e n t a l p h a - c e l l u l o s e , among 308 l o b l o l l y p i n e t r e e s . The c o r r e l a t i o n c o e f f i c i e n t , a l t h o u g h s i g n i f i c a n t , was o n l y 0.153, so t h a t t r a c h e i d l e n g t h e x p l a i n e d o n l y s l i g h t l y more t h a n two p e r c e n t o f t h e v a r i a t i o n i n c e l l u - l o s e c o n t e n t . E v e n t h o u g h i t i s a p p a r e n t t h a t t r a c h e i d l e n g t h and g r o s s wood c a r b o h y d r a t e c o n t e n t a r e r e l a t e d w i t h i n and between stems o f a s p e c i e s , t h e r e l a t i o n s h i p between s p e c i e s and i t s c a u s e , i s an a r e a o f r e l a t i v e l y l i t t l e i n f o r m a t i o n . M o r e o v e r , i t may be a p p r e c i a t e d t h a t c o m p a r i s o n s between g r o s s c e l l u l o s e c o n t e n t and s i n g l e c e l l c h a r a c t e r i s t i c s , s u c h as l e n g t h , a r e weak s i n c e t h e f o r m e r i s b a s e d on t h e e s t i m a t e o f numerous c e l l s i n a g i v e n volume o r w e i g h t o f wood, and t h e l a t t e r i s m e a s u r e d f o r i n d i v i d u a l c e l l s . T h e r e f o r e , s u c h c o m p a r i s o n s become l e s s s e n s i t i v e and t h i s may e x p l a i n why some o f t h e a u t h o r s f a i l e d t o show more t h a n two p e r c e n t v a r i a b i l i t y i n c e l l u l o s e c o n t e n t b a s e d on c e l l l e n g t h . N e v e r t h e l e s s , t h e s e f i n d i n g s do p r o v i d e e a r l i e r c l u e s f o r r e l a t i o n s h i p b e t w e e n l e n g t h a n d c a r b o h y d r a t e c o n t e n t i n wood. The r e s u l t s o b t a i n e d i n t h e p r e s e n t s t u d y ( F i g u r e s ' , : l t o 9) do show t h a t , i r r e s p e c t i v e o f s p e c i e s , t h e p o s s i b i l i t y e x i s t s f o r a s t r o n g r e l a t i o n s h i p b e t w e e n l e n g t h o f an i n d i v i - d u a l c e l l and w e i g h t o f i t s c a r b o h y d r a t e s k e l e t o n . H o w e v e r , as n o t e d b y some e a r l i e r w o r k e r s , i t i s n o t a p p a r e n t a t t h i s s t a g e w h e t h e r t h e r e l a t i o n s h i p o b t a i n e d i s due t o c h a n c e , i . e . , t h a t b o t h i n c r e a s e w i t h a g e , o r w h e t h e r t h e r e i s a common f u n d a m e n t a l c a u s e f o r s u c h a r e l a t i o n s h i p b e t w e e n s p e c i e s . T h e s e a nd r e l a t e d a s p e c t s w i l l be c o n s i d e r e d . 2. V a r i a t i o n b e t w e e n s p e c i e s As n o t e d , t h e p r i m a r y o b j e c t i v e o f t h i s i n v e s t i g a t i o n wa's t o r e l a t e t r a c h e i d w e i g h t t o i t s l e n g t h i n c o n i f e r s . I n F i g u r e 1 0 , a d e f i n i t e r e l a t i o n s h i p i s shown when c o m b i n e d d a t a f r o m a l l t h e n i n e c o n i f e r o u s woods ( e x c l u d i n g t h o s e f r o m com- p r e s s i o n wood a nd wood f o r m e d by f e r t i l i z e d t r e e s ) a r e p l o t t e d . The c o r r e l a t i o n s o b t a i n e d f r o m t h e c u r v i l i n e a r r e l a t i o n s h i p s a r e , h i g h l y s i g n i f i c a n t and a c c o u n t e d f o r 91.9 and 95.7 p e r c e n t o f t h e v a r i a b i l i t y i n h o l o c e l l u l o s e a n d a l p h a - c e l l u l o s e s k e l e t o n w e i g h t s , r e s p e c t i v e l y , by t h e l e n g t h f a c t o r a l o n e . When a c o n d i t i o n e d r e g r e s s i o n m o d e l (where t h e i n t e r c e p t i s z e r o ) was f i t t e d ( s e e F i g u r e 1 1 ) , t h e v a r i a b i l i t y a c c o u n t e d f o r was t h e same (91.9 p e r c e n t ) f o r h o l o c e l l u l o s e s k e l e t o n w e i g h t s , w h e r e a s " i t was s l i g h t l y l e s s (95.2) f o r a l p h a - c e l l u l o s e . I n s p i t e o f t h e s t r o n g r e l a t i o n s h i p s o b t a i n e d , i n d i - v i d u a l s p e c i e s d i f f e r e n c e s w e r e s i g n i f i c a n t . S u c h d i f f e r e n c e s c o u l d be e x p e c t e d s i n c e , f o r e a c h o f t h e s p e c i e s e x a m i n e d , t r a c h e i d c h a r a c t e r i s t i c s a r e a l s o d i f f e r e n t w i t h r e s p e c t t o d i a m e t e r and w a l l t h i c k n e s s ( 1 0 1 , 102, 1 1 6 ) . T h u s , d e s p i t e a s t r o n g r e l a t i o n s h i p b e t w e e n l e n g t h and w e i g h t i n e a c h s p e c i e s , some o f t h e s e d i f f e r e d s t a t i s t i c a l l y f r o m one a n o t h e r , i n r e s - p e c t i v e s l o p e s a n d l e v e l s . o f t h e r e g r e s s i o n l i n e s . A summary o f t h e s e s e p a r a t e a n a l y s e s i s g i v e n i n T a b l e s V and V I , f o r h o l o c e l l u l o s e and a l p h a - c e l l u l o s e t r a c h e i d s k e l e t o n s , r e s p e c - t i v e l y . A n a l y s e s f o r h o l o c e l l u l o s e t r a c h e i d s k e l e t o n s ( T a b l e V) . . i n d i c a t e t h a t , i n s p i t e o f t h e d i f f e r e n c e s , t h e r e i s s i m i l - a r i t y i n r e g r e s s i o n l i n e s l o p e s b e t w e e n s p e c i e s b e l o n g i n g t o one f a m i l y . Thus P i n u s l a m b e r t i a n a , P i c e a s i t c h e n s i s and P s e u d o t s u g a m e n z i e s i i , b e l o n g i n g t o t h e f a m i l y P i n a c e a e , d i d n o t d i f f e r f r o m one a n o t h e r i n s l o p e s ; i n a d d i t i o n , P i n u s l a m b e r t i a n a a n d P i c e a s i t c h e n s i s showed no d i f f e r e n c e i n t h e l e v e l s o f t h e same r e g r e s - s i o n l i n e s . S i m i l a r l y , T a x u s a nd C e p h a l o t a x u s , w h i c h a r e c l o s e l y r e l a t e d ( 1 5 9 ) , d i d n o t d i f f e r i n e i t h e r s l o p e s o r l e v e l s . I t i s o f i n t e r e s t t o n o t e t h e s i m i l a r i t y i n s l o p e s b e t w e e n P o d o c a r p u s d a c r y d i o d e s a n d P s e u d o t s u g a m e n z i e s i i , as w e l l as s i m i l a r i t y i n s l o p e s f o r t h e l a t t e r s p e c i e s t o g e t h e r w i t h T a x u s , J u n i p e r u s and C e p h a l o t a x u s . . The b e h a v i o u r o f a l p h a - c e l l u l o s e t r a c h e i d s k e l e t o n s , w i t h some e x c e p t i o n s , i s l i k e t h a t o f h o l o c e l l u l o s e , i n s h o w i n g s i m i l a r i t i e s b e t w e e n s p e c i e s b e l o n g i n g t o t h e same f a m i l y o r r e l a t e d f a m i l i e s ( T a b l e V I ) . S o m e t i m e s , h o w e v e r , t h e s i m i l a r i - t i e s n o t e d w e r e i n s p e c i e s b e l o n g i n g t o d i f f e r e n t f a m i l i e s , b u t o f t h e same t r a c h e i d l e n g t h c l a s s . A g a i n , P i n u s l a m b e r t i a n a a n d P i c e a s i t c h e n s i s d i d n o t d i f f e r f r o m one a n o t h e r i n s l o p e s . S i m i l a r l y , T a x u s a nd C e p h a l o t a x u s showed t h e same b e h a v i o u r w i t h r e g a r d t o s l o p e s , and t h e s e t w o , t o g e t h e r w i t h J u n i p e r u s , d i d n o t d i f f e r s i g n i f i c a n t l y . i n e i t h e r s l o p e s o r l e v e l s . I n t e r e s - t i n g l y , a l t h o u g h P s e u d o t s u g a d i d n o t d i f f e r f r o m P i n u s a n d P i c e a o f t h e same f a m i l y , i n s l o p e s o f r e g r e s s i o n l i n e s f o r h o l o - c e l l u l o s e s k e l e t o n s , i t d i f f e r e d s i g n i f i c a n t l y (5% l e v e l ) f r o m t h e s e two i n a l p h a - c e l l u l o s e c h a r a c t e r i s t i c s . A n o t h e r p o i n t o f i n t e r e s t i s t h e s i m i l a r i t y i n s l o p e s b e t w e e n A r a u c a r i a , S e q u o i a , P i n u s a n d P i c e a , as w e l l as t h a t b e t w e e n A r a u c a r i a a n d and P o d o c a r p u s . I n a d d i t i o n , A r a u c a r i a and S e q u o i a d i d riot-differ i n l e v e l s . The s l i g h t l y d i f f e r e n t b e h a v i o u r p a t t e r n o f a l p h a - c e l l u l o s e s k e l e t o n s , f r o m t h o s e o f h o l o c e l l u l o s e , c o u l d p o s s i b l y be r e l a t e d t o t h e s h r i n k a g e o f t h e t r a c h e i d s i n t h e r e m o v a l o f h e m i c e l l u l o s e s . P o s s i b l y t h e r a t i o o f s h r i n k a g e may be d i f - f e r e n t i n d i f f e r e n t s p e c i e s . T h e r e f o r e , i t w o u l d be o f i n t e r e s t t o p u r s u e t h i s m a t t e r f u r t h e r i n o r d e r t o d e t e r m i n e an e x a c t r e l a t i o n s h i p b e t w e e n s h r i n k a g e a n d a l k a l i t r e a t m e n t i n t h e s e f i b r e s k e l e t o n s , f o r d i f f e r e n t s p e c i e s , i n c l u d i n g - e a r l y w o o d a n d l a t e w o o d . 3. V a r i a t i o n w i t h i n c r e a s i n g age The s t r o n g i n d i v i d u a l s p e c i e s r e l a t i o n s h i p s b e t w e e n t r a c h e i d l e n g t h and w e i g h t , as w e l l as t h a t f o r a l l t h e s p e c i e s c o n s i d e r e d t o g e t h e r , has a l r e a d y b e e n p o i n t e d o u t . I t i s known, h o w e v e r , t h a t f o r any p a r t i c u l a r a n a t o m i c a l o r p h y s i c a l p r o p e r t y , g r e a t d i f f e r e n c e s c a n o c c u r w i t h i n a s i n g l e s t e m . I n v i e w o f t h i s , v a r i a t i o n o f t r a c h e i d w e i g h t and l e n g t h a c r o s s t h e r a d i u s was s t u d i e d . A v e r a g e t r a c h e i d w e i g h t and l e n g t h d a t a a r e l i s t e d i n T a b l e s I I I and I V f o r two D o u g l a s - f i r t r e e s . R a d i a l t r a c h e i d w e i g h t p a t t e r n s i n t h e b u t t l o g o f t h e 5 0 0 - y e a r - o l d t r e e a r e shown i n F i g u r e 1 2 . F o r c o m p a r i s o n , mean v a l u e s f o r t r a c h e i d l e n g t h o b t a i n e d f r o m m a c e r a t e d wood o f c e r t a i n i n c r e m e n t s , and s p e c i f i c g r a v i t y v a l u e s t a k e n f r o m K ennedy a n d W a r r e n (70) f o r t h i s wedge s a m p l e , w e r e a l s o i n c l u d e d i n t h e same f i g u r e . I t i s s e e n t h a t f o r b o t h e a r l y w o o d and l a t e w o o d t h e r e i s a m a r k e d i n c r e a s e i n t r a c h e i d w e i g h t up t o 80 y e a r s and a d e c r e a s e a f t e r a b o u t 150 y e a r s . T r a c h e i d l e n g t h f o r t h i s t r e e ( m a c e r a t e d t r a c h e i d s ) , as w e l l as t h e a v e r a g e t r a c h e i d l e n g t h o f . t h e s a m p l e s w e i g h e d ( T a b l e s I I I and I V ) , f o l l o w e d t h e same g e n e r a l t r e n d . I t i s o f i n t e r e s t t o n o t e t h a t t h e r a d i a l (wood) d e n - s i t y p a t t e r n d e s c r i b e d by Kennedy a n d W a r r e n (70) i s s i m i l a r t o t h e t r a c h e i d w e i g h t - l e n g t h p a t t e r n o b t a i n e d i n t h e p r e s e n t s t u d y , i n d i c a t i n g some i n t e r - r e l a t i o n s h i p b e t w e e n i n d i v i d u a l t r a c h e i d c h a r a c t e r i s t i c s and g r o s s wood s p e c i f i c g r a v i t y . The 56. l i t e r a t u r e d o e s n o t show t h i s c l e a r l y ( 4 7 ) . I n d i r e c t e v i d e n c e i s a v a i l a b l e , h o w e v e r , f r o m c e l l m o r p h o l o g i c a l s t u d i e s . I t i s p o s s i b l e t h a t w i t h i n c r e a s e i n l e n g t h t h e r e may be an i n c r e a s e i n p e r c e n t a g e o f c e l l w a l l as a p o r t i o n o f t o t a l c e l l v o l u m e , w h i c h i s c o m p a r a t i v e l y g r e a t e r f o r l o n g e r f i b r e s ; i . e . , c e l l w a l l v o l u m e p e r u n i t l e n g t h w o u l d be h i g h e r f o r l o n g e r t r a c h e i d s . I t i s r e a s o n a b l e , t h e r e f o r e , t o assume t h a t t r a c h e i d l e n g t h i s c l o s e l y c o r r e l a t e d w i t h wood d e n s i t y w i t h i n t h i s t r e e . T h u s , v a n B u i j t e n e n ( 1 3 8 ) , f r o m h i s s t u d y on s l a s h p i n e , c o n c l u d e d : T h e r e a p p e a r t o be two c o m p l e x e s o f p r o p e r - t i e s w h i c h a r e r a t h e r s t r o n g l y c o r r e l a t e d . They a r e s m a l l f i b e r d i a m e t e r , t h i n w a l l s , a l o w sum- merwood p e r c e n t a g e , s h o r t f i b e r s , and a l o w s p e c i - f i c g r a v i t y on t h e one h a n d , and l a r g e f i b e r d i a m e t e r , t h i c k w a l l s , a h i g h summerwood p e r c e n - t a g e , l o n g f i b e r s a n d a h i g h s p e c i f i c g r a v i t y on t h e o t h e r h a n d . T h e s e two c o m p l e x e s a r e t y p i c a l l y t h e j u v e n i l e p r o p e r t i e s v e r s u s t h e m a t u r e p r o p e r - t i e s , a l t h o u g h a p p a r e n t l y e v e n i n m a t u r e wood t h e s e p r o p e r t i e s seem t o be s t r o n g l y a s s o c i a t e d w i t h e a c h o t h e r . T h i s a s s o c i a t i o n , h o w e v e r , i s n o t r i g i d and i t i s p o s s i b l e t o f i n d a v a r i e t y o f c o m b i n a t i o n s . One c a n , f o r i n s t a n c e , f i n d wood s a m p l e s w i t h s m a l l f i b e r d i a m e t e r , t h i n w a l l s , a h i g h summerwood p e r c e n t a g e , and an a v e r a g e s p e c i f i c g r a v i t y . The r e a s o n s f o r a r e l a t i o n s h i p b e t w e e n t r a c h e i d l e n g t h and s p e c i f i c g r a v i t y w i t h i n t h i s s p e c i e s c o u l d a l s o be a t t r i b u t e d t o t h e i n c r e a s e i n amount o f c a r b o h y d r a t e f r a c t i o n i n t h e t r a c h e i d w h i c h , as shown a b o v e , i s h i g h l y c o r r e l a t e d w i t h t r a c - h e i d l e n g t h . N o t much i s known o n t h i s a s p e c t . Z o b e l and M c E l w e e (171) f o u n d a p o o r r e l a t i o n s h i p b e t w e e n c e l l u l o s e y i e l d and s p e c i f i c g r a v i t y on a w e i g h t b a s i s , b u t a v e r y s t r o n g 57. r e l a t i o n s h i p on a v o l u m e b a s i s i . e . , "a g i v e n v o l u m e o f h i g h s p e c i f i c g r a v i t y wood g i v i n g h i g h e r y i e l d s o f c e l l u l o s e t h a n t h e same v o l u m e o f l o w s p e c i f i c g r a v i t y -wood." S e v e r a l s t u d i e s i n d i c a t e h i g h l y s i g n i f i c a n t r e l a t i o n s h i p ; ' , b e t w e e n g r o s s wood c e l l u l o s e c o n t e n t and f i b r e l e n g t h (28, 6 0 ) , as w e l l as f i b r e l e n g t h and d e n s i t y ( 9 9 ) . A l s o , i t was n o t e d t h a t t h e p a t t e r n o f h o l o c e l l u l o s e v a r i a t i o n a c r o s s a g r o w t h i n c r e m e n t i s s i m i l a r t o t h a t o f t e n s i l e s t r e n g t h w h i c h i s h i g h l y s i g n i f i c a n t l y c o r - r e l a t e d ( r =,0.92) w i t h s p e c i f i c g r a v i t y . (156, 1 5 7 ) . I t i s p o s s i b l e t h a t s p e c i f i c g r a v i t y i s t h u s r e l a t e d i n d i r e c t l y t o c e l l u l o s e c o n t e n t i n t h e t r a c h e i d s . Wood f o r m e d i n t h e r e g i o n n e a r t h e p i t h i s c a l l e d j u v e n i l e o r c o r e w o o d ; when c o m p a r e d t o m a t u r e wood i t h a s d i f f e r e n t p r o p e r t i e s s u c h a s - s h o r t e r t r a c h e i d s and l o w e r c e l l u - l o s e p e r c e n t a g e , s p e c i f i c g r a v i t y , p e r c e n t a g e o f l a t e w o o d , a nd t e n s i l e s t r e n g t h ( 4 1 ) . D a t a f r o m t h i s s t u d y a l s o show t h a t w e i g h t s o f t h e i n d i v i d u a l t r a c h e i d s k e l e t o n s f r o m j u v e n i l e w o o d 1 ( D o u g l a s - f i r I n c r e m e n t 1 0 ) , a p p a r e n t l y , a r e w e l l b e l o w t h e l e v e l f o u n d i n m a t u r e and o v e r m a t u r e wood i n c r e m e n t s ( F i g u r e 1 2 ) . S t a t i s t i c a l a n a l y s e s , h o w e v e r , d i d n o t c o n f i r m t h i s e n t i r e l y ( s e e T a b l e s V I I and V I I I ) . A n a l y s e s f o r h o l o - c e l l u l o s e s k e l e t o n s i n d i c a t e t h a t no s i g n i f i c a n t d i f f e r e n c e i n "'"Only I n c r e m e n t 10 i s c o n s i d e r e d as j u v e n i l e wood, s i n c e McKimmy (94) showed f o r D o u g l a s - f i r t h a t t h e j u v e n i l e p e r i o d e x t e n d s t o a b o u t 10 a n n u a l i n c r e m e n t s f r o m t h e p i t h . A l s o , t r a c h e i d l e n g t h and s p e c i f i c g r a v i t y d a t a f o r t h i s t r e e do n o t w a r r a n t c l a s s i f y i n g I n c r e m e n t 20 as j u v e n i l e wood. 58. w e i g h t c o u l d be f o u n d between j u v e n i l e wood t r a c h e i d s as compared w i t h t h o s e f r o m m a t u r e and o v e r m a t u r e wood t r a c h e i d s . C o n t r a r y t o t h i s , a n a l y s e s f o r a l p h a - c e l l u l o s e s k e l e t o n s ( T a b l e V I I I ) i n d i c a t e t h a t , f o r t h e same t r a c h e i d l e n g t h , t h e j u v e n i l e wood t r a c h e i d s a r e s i g n i f i c a n t l y l o w e r i n w e i g h t com- p a r e d t o m a t u r e (A) and o v e r m a t u r e wood (B) t r a c h e i d s . The o b v i o u s r e a s o n f o r t h e s e a p p a r e n t l y c o n f l i c t i n g c o n c l u s i o n s i s t h a t q u a l i t a t i v e d i f f e r e n c e s may e x i s t ; i . e . , f o r t h e same t r a c h e i d l e n g t h , j u v e n i l e wood t r a c h e i d s w i l l h a v e more hemi- c e l l u l o s e f r a c t i o n t h a n t h o s e f r o m m a t u r e and o v e r m a t u r e wood. L a r s o n (78) has n o t e d some v e r y d e f i n i t e q u a l i t a t i v e c h a n g e s i n w a l l c h e m i c a l c o m p o s i t i o n w i t h i n c r e a s e i n age f o r r e d p i n e . From t h i s and o t h e r c o n s i d e r a t i o n s , he c o n c l u d e d t h a t t h e s e c h a n g es a p p e a r t o be i n h e r e n t and may by t y p i f i e d by t h e . t r e n d e x p r e s s e d i n t h e e a r l y w o o d . Some i n d i r e c t s u p p o r t c o u l d a l s o be f o u n d f r o m Z o b e l and McElwee's (171) work on l o b l o l l y p i n e . They f o u n d t h a t , b a s e d on d r y w e i g h t , y i e l d s f r o m m a t u r e wood r a n 3.5 p e r c e n t h i g h e r f o r w a t e r - r e s i s t a n t c a r b o h y d r a t e (WRC), whereas t h o s e f o r a l p h a - c e l l u l o s e were 7.5 p e r c e n t h i g h e r when compared t o t h e y i e l d f r o m j u v e n i l e wood from-the same t r e e s . As p o i n t e d o u t by L a r s o n ( 7 6 ) , f o r m a t i o n o f j u v e n i l e wood i s a s s o c i a t e d w i t h t h e p r o l o n g e d i n f l u e n c e o f t h e a p i c a l m e r i s t e m , d u r i n g t h e g r o w i n g s e a s o n , i n t h e r e g i o n o f t h e a c t i v e crown. As t h e crown moves upward i n t h e o l d e r t r e e , t h e cambium a t a g i v e n h e i g h t becomes l e s s s u b j e c t t o t h e d i r e c t i n f l u e n c e s o f t h e e l o n g a t i n g crown r e g i o n and a d u l t wood i s f o r m e d ( 1 1 1 ) . T h e r e f o r e , t h e r e a s o n f o r t h e s i g n i f i c a n t l y l o w e r w e i g h t o f a l p h a - c e l l u l o s e s k e l e t o n s i n j u v e n i l e wood i s e v i d e n t , s i n c e wood p r o d u c e d d u r i n g t h e j u v e n i l e p e r i o d i s o f d i f f e r e n t q u a l i t y as s t a t e d a b ove, w h i c h q u a l i t y w i l l be r e f l e c t e d i n t h e i n d i v i d u a l c e l l . S e v e r a l arguments h a v e b e e n o f f e r e d t o e x p l a i n t h e d e c r e a s e i n c e l l u l o s e p e r c e n t a g e and s p e c i f i c g r a v i t y i n o v e r m a t u r e wood ( s e e T a b l e s V I I ( C ) and V I I I ( C ) ; and F i g u r e 1 2 ) . F o r example, H a l e and C l e r m o n t (52) e x p l a i n e d t h a t o v e r - m a t u r e wood ( t h a t f o r m e d i n t h e o u t e r p o r t i o n s ) o f r e l a t i v e l y o l d t r e e s (300 t o 400 y e a r s ) has c h a r a c t e r i s t i c a l l y t h i n - w a l l e d p r o s e n c h y m a . w i t h h i g h e r l i g n i n and l o w e r a l p h a - c e l l u l o s e p e r - c e n t a g e t h a n wood f o r m e d d u r i n g e a r l i e r s t a g e s o f g r o w t h . Kennedy and W a r r e n (70) s u p p o r t e d D o e r n e r 1 s c o n c e p t (3 5) i n w h i c h he e x p l a i n s t h e i n c r e a s e and s u b s e q u e n t d e c r e a s e i n s p e c i f i c g r a v i t y on m e c h a n i s t i c p r i n c i p l e s . D o e r n e r s u g g e s t s t h a t when t h e t r e e i s g r o w i n g a p p r e c i a b l y i n h e i g h t , b e n d i n g s t r e s s e s a r e g r e a t e r , t h e r e b y n e c e s s i t a t i n g t h e , f o r m a t i o n o f de n s e p e r i p h e r a l wood t o augment a r e l a t i v e l y s m a l l s e c t i o n m o d u l u s . A t a d v a n c e d a g e s , when t h e r a t e o f v e r t i c a l t o r a d i a l i n c r e m e n t has d e c r e a s e d m a r k e d l y , t h e t r e e c o n t i n u e s t o r e a c t e f f i c i e n t l y t o t h e r e d u c e d s t r e s s s t i m u l u s by p r o d u c i n g l e s s - d e n s e wood t o c o u n t e r a c t t h e d e v e l o p m e n t o f an e x c e s s i v e l y l a r g e 60. s e c t i o n m o d u l u s . F o r b o t h c a s e s he s u g g e s t s t h a t t h e d e n s i t y c h a n g e s a r e p r o d u c e d by t h e r e a c t i o n s o f d e v e l o p i n g t i s s u e s t o t h e s t i m u l i . The f o r m e r a u t h o r s (52) t h u s s u p p o r t a d i r e c t age e f f e c t i n d e p e n d e n t o f t r e e d i a m e t e r , w h e r e a s D o e r n e r i n f e r s an i n d i r e c t one s i n c e d i a m e t e r i n c r e a s e s w i t h a g e . R e s u l t s o f t h e p r e s e n t s t u d y on s i n g l e t r a c h e i d w e i g h t s a r e i n a g r e e m e n t w i t h t h e a b o v e c o n c l u s i o n s b a s e d on g r o s s wood a n a l y s e s , and p r o v i d e a f u r t h e r p o i n t o f p r o o f i n t h a t c h a n g e s i n g r o w t h b e h a v i o u r o f t r e e s a r e r e f l e c t e d more d i r e c t l y i n t h e d e v e l o p m e n t o f t h e i n d i v i d u a l c e l l w h e t h e r i n n o r m a l o r r e a c t i o n wood f o r m a t i o n ( s e e b e l o w ) . A d d i t i o n a l c o n f i r m a t i o n was o b t a i n e d by t h e a n a l y s i s o f c o v a r i a n c e t e s t ( T a b l e s V I I a n d V I I I ) i n w h i c h t r a c h e i d w e i g h t s o f m a t u r e wood ( I n c r e m e n t s No. 20, 80, 150) w e r e c o m p a r e d w i t h t h o s e o f o v e r - m a t u r e wood ( I n c r e m e n t s No. 3 0 0 , 4 0 0 ) , a f t e r a d j u s t i n g f o r d i f f e r e n c e s i n t r a c h e i d l e n g t h . The s i g n i f i c a n t 'F' v a l u e s f o r ' s l o p e s ' i n b o t h c a s e s i n d i c a t e t h a t t h e s e p o p u l a t i o n s d i f f e r . T h i s means t h a t t h e s i g n i f i c a n t l y l o w e r t r a c h e i d w e i g h t s i n o v e r m a t u r e wood c o u l d be a t t r i b u t e d t o some f a c t o r s u c h as t h i n w a l l e d p r o s e n c h y m a c e l l s as s u g g e s t e d by H a l e and C l e r m o n t ( 5 2 ) . 4. V a r i a t i o n b e t w e e n r e g u l a r ( n o r m a l ) and c o m p r e s s i o n wood R e a c t i o n wood o f c o n i f e r s , u s u a l l y r e f e r r e d t o as com- p r e s s i o n wood, i s d e v e l o p e d as p a r t o f an o r i e n t a t i o n m e c h a n i s m , i . e . , a g e o t r o p i c r e a c t i o n t o an i n e r t i a l f o r c e ( 1 5 1 ) . As s t a t e d by L a r s o n ( 8 0 ) : The i n t e r a c t i o n between a u x i n and s u c r o s e i n p r o d u c i n g c o m p r e s s i o n wood r e s u l t s i n t r a c - h e i d s w i t h l a r g e r d i a m e t e r s and e x t r e m e l y t h i c k c e l l w a l l s . P h y s i c a l l y , c o m p r e s s i o n wood t r a c - h e i d s a r e somewhat s i m i l a r t o t h o s e o f j u v e n i l e wood p r o d u c e d i n t h e h i g h a u x i n e n v i r o n m e n t i n c l o s e p r o x i m i t y t o t h e crown. C h e m i c a l l y , t h e n o r m a l m e t a b o l i c pathways a p p e a r t o be a l t e r e d d u r i n g c o m p r e s s i o n wood f o r m a t i o n s o t h a t p r o - d u c t i o n o f - c o n s t i t u e n t s n o r m a l l y c o n f i n e d t o t h e o u t e r w a l l l a y e r s i s p e r p e t u a t e d a c r o s s t h e w a l l . Wardrop and D a d s w e l l (146) p o i n t e d o u t t h a t t h e r e d u c t i o n i n t r a c h e i d l e n g t h ( C h a p t e r I I ) r e s u l t s f r o m an i n c r e a s e i n " t h e number o f a n t i c l i n a l d i v i s i o n s i n t h e cambium a s s o c i a t e d w i t h t h e r a p i d e c c e n t r i c r a d i a l g r o w t h t h a t i s u s u a l w i t h t h e d e v e l o p m e n t o f c o m p r e s s i o n wood. Thus t h e r e a p p e a r s t o be a s i m i l a r i t y b etween s u c h c e l l s and t h e t r a c h e i d s f o r m e d d u r i n g t h e y o u t h f u l p e r i o d o f t h e l i f e o f t h e t r e e . I n many i n v e s t i g a t i o n s , c o m p a r i s o n s h a ve been f r e q u e n - t l y made w i t h m a t e r i a l f r o m t h e q u a d r a n t a d j a c e n t t o t h e com- p r e s s i o n wood t o t y p i f y n o r m a l wood. I n t h i s s t u d y , however, c o m p a r i s o n s a r e made on t h e b a s i s o f same t r a c h e i d l e n g t h , s i n c e t h e y may be more m e a n i n g f u l and s e n s i t i v e . M o r e o v e r , i n s p i t e o f t h e o b v i o u s m o r p h o l o g i c a l d i f f e r e n c e s , t h e ma i n d i f f e r e n c e between c o m p r e s s i o n wood and n o r m a l wood i s one o f d e g r e e o n l y ( m i l d e s t t o e x t r e m e t y p e ) , o t h e r d e v e l o p m e n t a l f e a t u r e s b e i n g s i m i l a r t o t h o s e o c c u r r i n g i n n o r m a l wood ( s e e 1 4 6 ) . A v e r a g e v a l u e s o f c o m p r e s s i o n wood t r a c h e i d w e i g h t s and t h e i r mean l e n g t h s a r e c o n t a i n e d i n T a b l e s I and I I , f o r h o l o - and a l p h a - c e l l u l o s e s k e l e t o n s , r e s p e c t i v e l y . R e l a t i o n - s h i p s b e t w e e n t r a c h e i d l e n g t h and w e i g h t a r e shown i n F i g u r e 13. I t i s a p p a r e n t t h a t , e v e n i n t h e f o r m a t i o n o f c o m p r e s s i o n wood, t h e r e i s a g a i n a h i g h l y s i g n i f i c a n t c o r r e l a t i o n between t r a c h e i d w e i g h t and l e n g t h , e x p l a i n i n g 71.7 and 90.0 p e r c e n t v a r i a t i o n i n h o l o c e l l u l o s e and a l p h a - c e l l u l o s e s k e l e t o n w e i g h t s , r e s p e c t i v e l y , by t h e l e n g t h f a c t o r a l o n e . T h i s b e h a v i o u r o f c o m p r e s s i o n wood r e a f f i r m s t h e p r e v i o u s l y e s t a b l i s h e d r e l a t i o n - s h i p f o r n o r m a l wood ( s e e F i g u r e ,9) . A g e n e r a l c o m p a r i s o n between c o m p r e s s i o n wood and n o r m a l wood i n d i c a t e s t h a t , f o r t h e same t r a c h e i d l e n g t h , no s i g n i f i c a n t d i f f e r e n c e c a n be f o u n d w i t h r e s p e c t t o h o l o c e l l u - l o s e t r a c h e i d s k e l e t o n s ( F i g u r e 1 3 ) . I n c o n t r a s t , d i f f e r e n c e s seem t o e x i s t f o r a l p h a - c e l l u l o s e s k e l e t o n w e i g h t s , s u g g e s t i n g t h e e x i s t e n c e o f q u a l i t a t i v e a n d / o r q u a n t i t a t i v e d i f f e r e n c e s . E x a m i n a t i o n o f t h e r e l a t i o n s h i p s g i v e n i n F i g u r e 13 as w e l l as t h e o r i g i n a l d a t a , however, show t h a t , f o r t h e same t r a c h e i d l e n g t h , c o m p r e s s i o n wood t r a c h e i d s may have s l i g h t l y h i g h e r a l p h a - c e l l u l o s e t h a n j u v e n i l e wood (where t r a c h e i d l e n g t h s were m a i n l y u n d e r 2.3 0 mm) b u t a l o w e r amount compared t o m a t u r e wood t r a c h e i d s (where t r a c h e i d l e n g t h s were m o s t l y , o v e r 2.30 mm). I n a d d i t i o n , s i n c e c o m p r e s s i o n wood t r a c h e i d s d i d n o t d i f f e r f r o m t h o s e o f n o r m a l wood i n h o l o c e l l u l o s e c o n t e n t , t h e y must c o n t a i n more h e m i c e l l u l o s e f r a c t i o n t h a n c o m p a r a b l e n o r m a l wood t r a c h e i d s , i . e . , t h e d i f f e r e n c e between h o l o - and a l p h a - c e l l u l o s e . The above b e h a v i o u r c o u l d be r e l a t e d t o a g e - d e p e n d e n t i n f l u e n c e . F o r example, i n t h e p r e s e n t i n s t a n c e , t h e compres-. s i o n wood s e l e c t e d was f r o m I n c r e m e n t s 73 and 83 f r o m t h e p i t h ( A p p e n d i x I ) . W i t h t h i s b a c k g r o u n d , i t c a n be p r o p o s e d f r o m t h e d a t a o b t a i n e d t h a t , d e s p i t e a d e c r e a s e i n t r a c h e i d l e n g t h a s s o c i a t e d w i t h c o m p r e s s i o n wood f o r m a t i o n , t h e age e f f e c t i s s t i l l s u p e r i m p o s e d i n t h e c a m b i a l b e h a v i o u r . Hence t r a c h e i d s a r e p r o d u c e d w i t h s l i g h t l y l o w e r amount o f a l p h a - c e l l u l o s e t h a n c o m p a r a b l e n o r m a l wood t r a c h e i d s . S i m i l a r l y , t h e s l i g h t l y h i g h e r a l p h a - c e l l u l o s e v a l u e s as compared t o t h o s e o f j u v e n i l e wood t r a c h e i d s c o u l d be e x p l a i n e d on t h e same p r i n c i p l e . The r e s u l t s h e r e a r e c o n s i s t e n t w i t h t h e f i n d i n g s o f A u s t r a l i a n w o r k e r s ( 2 7 ) . T h e s e a u t h o r s e m p h a s i z e d t h a t a l l d e g r e e s o f c o m p r e s s i o n wood may be e n c o u n t e r e d f r o m t h e m i l d e s t f o r m t o t h e v e r y e x t r e m e t y p e s , and t h a t t h e r e i s no b a s i c d i f f e r e n c e between t h e s e and n o r m a l wood, i n t h e mechanism o f c e l l d i v i s i o n , i n t h e r e a d j u s t m e n t o f d a u g h t e r c e l l s , o r i n t h e p r o g r e s s i v e d e v e l o p m e n t o f c e l l w a l l , and t h e d i f f e r e n c e i s i n d e g r e e o n l y . S i n c e , i n t h e p r e s e n t s t u d y , c a r e was t a k e n t o i n c l u d e t h e ext r e m e c a s e o f c o m p r e s s i o n wood i t was p o s s i b l e t o b r i n g o u t t h e s e d i f f e r e n c e s o f d e g r e e . I n summary, i t c a n be s t a t e d t h a t t h e b a s i c r e l a t i o n s h i p , d e v e l o p e d w i t h n o r m a l wood t r a c h e i d s , seems t o h o l d f o r c o m p r e s s i o n wood as w e l l , i n d i c a t i n g a common r e g u l a t o r y mech- a n i s m i n c o n i f e r c a m b i a l m o r p h o l o g y . H o w e v e r , c o m p r e s s i o n wood t r a c h e i d s d i f f e r f r o m n o r m a l wood t r a c h e i d s i n s h o w i n g some q u a l i t i a t i v e d i f f e r e n c e s w i t h r e g a r d t o t h e b a s i c c a r b o h y d r a t e s k e l e t a l f r a c t i o n ( a l p h a - c e l l u l o s e ) . The r e m a r k a b l e a b i l i t y o f t h e t r e e f o r q u i c k r e a d j u s t m e n t s a t c e l l u l a r l e v e l , t o c h a n g e s b r o u g h t a b o u t by e x t r i n s i c f a c t o r s , i s c l e a r l y e m p h a s i z e d i n t h e p h y s i o l o g y o f c o m p r e s s i o n wood f o r m a t i o n . 5. V a r i a t i o n b e t w e e n e a r l y w o o d a n d l a t e w o o d I t h a s l o n g b e e n r e c o g n i z e d t h a t e a r l y w o o d a nd l a t e - wood o f c o n i f e r o u s w oods, w i t h i n a g r o w t h i n c r e m e n t , h a v e d i f f e r e n t m o r p h o l o g i c a l , p h y s i c a l a n d c h e m i c a l p r o p e r t i e s a n d b e h a v i o u r s . I n a l m o s t e v e r y r e s p e c t t h e d i f f e r e n c e s b e t w e e n t h e s e two t i s s u e t y p e s may be e s s e n t i a l l y as g r e a t a s , i f n o t g r e a t e r t h a n , t h o s e b e t w e e n wood z o n e s w i t h i n a s i n g l e s t e m , . wood f r o m two members o f a s i n g l e t r e e s p e c i e s , o r b e t w e e n two s p e c i e s ( 5 9 ) . However, a n a t o m i c a l e v i d e n c e seems t o i n d i c a t e no s i g n i f i c a n t d i f f e r e n c e i n t h e a c t u a l amounts o f c e l l w a l l m a t e r i a l p e r t r a c h e i d i n e a r l y w o o d a nd l a t e w o o d t r a c h e i d s i n c o n i f e r s ( 1 4 , 68, 1 2 1 a ) . The a p p a r e n t v a r i a t i o n s o b s e r v e d i n s p e c i f i c g r a v i t y a c r o s s an i n c r e m e n t h a v e b e e n a t t r i b u t e d l a r g e l y t o d i f f e r e n c e s i n t r a c h e i d v o l u m e r a t h e r t h a n w a l l s u b s t a n c e ( 7 0 ) . I t i s o f i n t e r e s t , t h e r e f o r e , t o e x a m i n e t h e v a r i a t i o n o f i n d i v i d u a l t r a c h e i d s f r o m t h e two a r b i t r a r y - f r a c t i o n s o f t h e i n c r e m e n t . As a p r e l i m i n a r y a p p r o a c h t o t h i s , t h e t r a c h e i d l e n g t h - w e i g h t r e l a t i o n s h i p s o b t a i n e d a r e r e - e x a m i n e d s e p a r a t e l y f o r e a r l y w o o d a nd l a t e w o o d . The c u r v i l i n e a r m u l t i p l e r e g r e s s i o n r e l a t i o n s h i p s o f b e s t f i t , f o r t h e c o m b i n e d d a t a o f t h e n i n e c o n i f e r o u s w oods, a r e g i v e n b e l o w . HOLOCELLULOSE ALPHA-CELLULOSE E a r l y w o o d E a r l y w o o d Y = 0.0291 + 0 . 0 5 0 6 X 2 Y = -0.2418 + 0.1845X + 0 . 0 3 7 7 X 2 2 r = 0.951; SEE = 0.198; R 2 = 0.972; SEE = 0.130; DF = 345 DF = 356 L a t e w o o d L a t e w o o d Y = -0.3770 + 0.2365X Y = -0.3705 + 0.2602X + 0 . 0 3 1 0 X 2 + 0 . 0 3 0 1 X 2 0.939; SEE = 0.263; R 2 = 0.959; SEE = 0.169; DF = 447 DF = 383 E a r l y w o o d + L a t e w o o d E a r l y w o o d + L a t e w o o d Y = -0.1708 + 0.1201X Y = -0.3019 + 0.2214X + 0 . 0 4 1 2 X 2 + 0 . 0 3 4 1 X 2 R 2 = 0.936; SEE = 0.255; R 2 = 0.965; SEE = 0.152; DF = 794 DF = 742 Where Y = o v e n d r y w e i g h t o f t r a c h e i d i n g x 10 X = t r a c h e i d l e n g t h i n mm - g r e e n c o n d i t i o n . T h e s e r e l a t i o n s h i p s c o n f i r m t h a t , i r r e s p e c t i v e o f o r i g i n o f t h e wood w i t h i n t h e i n c r e m e n t , t h e r e i s a h i g h . d e g r e e o f c o r r e l a t i o n between l e n g t h and w e i g h t among t r a c h e i d s , The 2 i n d i v i d u a l r v a l u e s obtained,, f o r e a r l y w o o d and l a t e w o o d d a t a , and t h e i r s t a n d a r d e r r o r s o f e s t i m a t e , d o n o t d i f f e r much f r o m t h o s e o f t h e combined v a l u e s . More d e t a i l e d a n a l y s e s were done o f b o t h h o l o - and a l p h a - c e l l u l o s e d a t a t o e x p l o r e f u r t h e r t h e b e h a v i o u r o f e a r l y w o o d and l a t e w o o d , as w e l l as t o d e t e r m i n e d i f f e r e n c e s b etween and w i t h i n s p e c i e s . R e s u l t s b a s e d on ' t ' t e s t s i n d i c a t e t h a t , f o r h o l o c u l l u l o s e , w e i g h t s o f l a t e w o o d t r a c h e i d s f r e q u e n t l y ( s i x o f t h e n i n e s p e c i e s ) a r e s i g n i f i c a n t l y h i g h e r t h a n t h o s e o f e a r l y w o o d t r a c h e i d s ( T a b l e I ) . T h i s t e n d e n c y i s l e s s p r o - n o u n c e d f o r a l p h a - c e l l u l o s e s k e l e t o n s , where f i v e o f t h e n i n e s p e c i e s e x amined d i d n o t show any s i g n i f i c a n t d i f f e r e n c e i n t r a c h e i d w e i g h t between t h e two t i s s u e f r a c t i o n s ( T a b l e I I ) . I n t e r e s t i n g l y , i n b o t h c a s e s ( h o l o - and a l p h a - c e l l u l o s e ) , ' t ' t e s t s f o r t h e combined d a t a showed t h a t l a t e w o o d t r a c h e i d s a r e a l w a y s s i g n i f i c a n t l y h i g h e r i n w e i g h t t h a n e a r l y w o o d t r a c - h e i d s , t h u s c o n f i r m i n g t h e f i n d i n g s o f o t h e r s b a s e d on g r o s s a n a l y s e s (1, 52, 1 1 4 ) . I t i s t o be n o t e d , however, t h a t where s i g n i f i c a n t d i f f e r e n c e s e x i s t , l a t e w o o d t r a c h e i d s a r e l o n g e r t h a n t h e i r c o u n t e r p a r t s i n e a r l y w o o d ( T a b l e s I and I I ) . B e c a u s e o f t h e above r e a s o n , a n a l y s e s o f c o v a r i a n c e t e s t s were done t o compare t r a c h e i d w e i g h t s a f t e r a d j u s t i n g f o r l e n g t h d i f f e r e n c e s . R e s u l t s o f t h e s e a n a l y s e s a r e g i v e n i n T a b l e s I X and X. I t c a n be s e e n t h a t , f o r h o l o c e l l u l o s e s k e l e t o n s , no s i g n i f i c a n t d i f f e r e n c e s c a n be f o u n d b e t w e e n t h e s i x t r e e s o f D o u g l a s - f i r e x a m i n e d ( T a b l e I X ) . As e x p e c t e d , d i f f e r e n c e s e x i s t b e t w e e n s p e c i e s ( T a b l e X) w h i c h , as a l r e a d y m e n t i o n e d , a r e a t t r i b u t a b l e t o d i f f e r e n c e s i n t h e , ' s l o p e s ' and ' l e v e l s ' ( T a b l e s V and V I ) . The p o i n t o f i n t e r e s t , h o w e v e r , i s t h e b e h a v i o u r o f e a r l y w o o d and l a t e w o o d , i n t h a t l a t e w o o d h o l o c e l l u l o s e t r a c h e i d s k e l e t o n s a r e s i g n i f i c a n t l y h e a v i e r t h a n e a r l y w o o d , b o t h w i t h i n a n d b e t w e e n s p e c i e s . I n c o n t r a s t t o t h e a b o v e , no s u c h d i f f e r e n c e s a r e e v i d e n t i n a l p h a - c e l l u l o s e s k e l e t o n w e i g h t s . T h i s i s n o t o n l y t r u e f o r t h e s i x D o u g l a s - f i r t r e e s e x a m i n e d , b u t a l s o f o r a l l t h e n i n e c o n i f e r o u s woods. I n r e g a r d t o o v e r a l l b e h a v i o u r , h i g h l y s i g n i f i c a n t d i f f e r e n c e s a r e f o u n d b e t w e e n s p e c i e s ( T a b l e X) w h i l e w i t h i n s p e c i e s ( i . e . , s i x D o u g l a s - f i r t r e e s ) , t h e d i f - f e r e n c e s a r e s i g n i f i c a n t o n l y a t t h e 5 p e r c e n t l e v e l ( T a b l e I X ) . The l a s t m e n t i o n e d d i f f e r e n c e s c o u l d w e l l r e s u l t f r o m t h e a l - r e a d y d e s c r i b e d c h a r a c t e r i s t i c s o f j u v e n i l e wood and c o m p r e s s i o n wood. I n s u m m a r i z i n g t h e v a r i a t i o n s i n e a r l y w o o d and l a t e w o o d t r a c h e i d s k e l e t o n w e i g h t s , f o r a l l t h e n i n e c o n i f e r o u s woods, i t i s e v i d e n t t h a t , f o r t h e same t r a c h e i d l e n g t h , e a r l y w o o d and l a t e w o o d w o u l d h a v e s i m i l a r amounts o f a l p h a - c e l l u l o s e , y e t d i f f e r i n g amounts o f h e m i c e l l u l o s e s . Q u a l i t a t i v e d i f f e r e n c e s 68. i n t h e h e m i c e l l u l o s e f r a c t i o n have a l s o b e en r e p o r t e d by o t h e r w o r k e r s f r o m g r o s s wood and t i s s u e a n a l y s e s (1, 78, 90, 114) b u t , i n a d d i t i o n , t h e s e r e p o r t s show d i f f e r e n c e s i n a l p h a - . c e l l u l o s e p e r c e n t a g e as w e l l . I n one s t u d y , however, b a s e d on t i s s u e a n a l y s i s on S c o t c h p i n e , M e i e r (90) o b t a i n e d a b o u t t h e same p e r c e n t a g e o f a l p h a - c e l l u l o s e f o r b o t h e a r l y w o o d and l a t e - wood. I n t h i s r e s p e c t t h e p r e s e n t r e s u l t s a r e i n agre e m e n t w i t h M e i e r ' s f i n d i n g s . As n o t e d , . g r o s s a n a l y s e s g i v e o n l y an e s t i m a t e o f t h e t o t a l f r a c t i o n ( p e r c e n t a g e ) o f c a r b o h y d r a t e f o r t h e t i s s u e as a w h o l e , w h i c h adds t o t h e d i f f i c u l t y o f i n t e r p r e t i n g d a t a b a s e d on i n d i v i d u a l c e l l w e i g h t s . T h a t i s , g r o s s a n a l y s e s a r e n o t s e n s i t i v e enough t o r e f l e c t t h e t r u e v a l u e s f o r w e i g h t s o f i n d i v i d u a l t r a c h e i d s f r o m d i f f e r e n t o r i g i n s . E v i d e n c e f o r a Common R e l a t i o n s h i p i n C o n i f e r s From f o r e g o i n g o b s e r v a t i o n s on t h e r e l a t i o n s h i p s b e tween l e n g t h o f t r a c h e i d and i t s c a r b o h y d r a t e f r a c t i o n , i t i s e v i d e n t t h a t a common r e l a t i o n s h i p c o u l d be d e r i v e d f o r a l l c o n i f e r s i r r e s p e c t i v e o f wood o r i g i n . I n o t h e r words, t h e r e l a t i o n s h i p i s n o t n e c e s s a r i l y an age d e p e n d e n t one i n t h a t b o t h t r a c h e i d l e n g t h and c e l l u l o s e c o n t e n t i n c r e a s e w i t h age i n a stem w i t h i n a s p e c i e s . As e x p e c t e d , t h e r e a r e s t a t i s t i c a l d i f f e r e n c e s i n t h e s l o p e s o f r e g r e s s i o n l i n e s d e p e n d i n g on s p e c i e s . I f , f r o m p r a c t i c a l r a t h e r t h a n s t r i c t l y a c a d e m i c s t a t i s t i c a l c o n s i d e r a t i o n s , a common r e g r e s s i o n l i n e c o u l d be f i t t e d f o r a l l t h e s e d a t a , i t w o u l d e x p l a i n a s i g n i f i c a n t p a r t o f t h e v a r i a b i l i t y i n t r a c h e i d w e i g h t , w i t h i n t o l e r a b l e l i m i t s , o f s t a n d a r d e r r o r o f t h e e s t i m a t e . S u c h a r a n d o m i - z a t i o n o f t r a c h e i d s w o u l d p r o v i d e a u n i q u e m e a s u r e f o r e x a m i - n i n g f u n d a m e n t a l m o r p h o l o g i c a l and c h e m i c a l b e h a v i o u r o f . t h e c o n i f e r wood b a s i c u n i t , s i n c e v a r i a b i l i t y i n t r o d u c e d by s p e c i e s , e n v i r o n m e n t a l a n d o n t o g e n e t i c f a c t o r s i s m i n i m i z e d . T h u s , a g e n e r a l b e h a v i o u r a l p a t t e r n c o u l d be d e d u c e d . The c o m b i n e d r e l a t i o n s h i p s o f F i g u r e 10 c o n f i r m t h a t , d e s p i t e s t a t i s t i c a l d i f f e r e n c e s i n s l o p e s b e t w e e n s p e c i e s , a s i g n i f i c a n t r e l a t i o n s h i p e x i s t s among t r a c h e i d s o f d i f f e r e n t c o n i f e r s , i n d i c a t i n g t h a t t h e p h y s i o l o g i c a l m e c h a n i s m s g o v e r n i n g t r a c h e i d l e n g t h a n d w e i g h t r e l a t i o n s h i p s a r e t h e same w i t h i n and b e t w e e n s p e c i e s , i n c l u d i n g wood f o r m e d o v e r a w i d e r a n g e o f t r e e a g e , n o r m a l wood and r e a c t i o n wood. T h u s , i f ' t r a c h e i d ' i s c o n s i d e r e d as one p o p u l a t i o n , t h e r e l a t i o n s h i p s shown i n F i g u r e s 10 and 11 a r e j u s t i f i e d , s i n c e t h e c o r r e l a t i o n s o b t a i n e d a r e h i g h l y s i g n i f i c a n t . I n a d d i t i o n , t h e s t a n d a r d e r r o r s o f e s t i m a t e s a r e no h i g h e r t h a n t h o s e n o t e d f o r some o f t h e i n d i - v i d u a l s p e c i e s ( e . g . , F i g u r e 9 - - H o l o c e l l u l o s e and F i g u r e 1-- A l p h a - c e l l u l o s e ) , i n d i c a t i n g t h e p a r a m o u n t i m p o r t a n c e o f l e n g t h f a c t o r w i t h r e g a r d t o t h e c e l l l e n g t h - w e i g h t r e l a t i o n s h i p s . I t i s e n t i r e l y p o s s i b l e t h a t a d d i n g t h e e f f e c t s o f o t h e r m o r p h o l o - g i c a l c h a r a c t e r i s t i c s , s u c h as c e l l d i a m e t e r . a n d w a l l t h i c k n e s s , n o t done h e r e i n , w o u l d i m p r o v e t h e r e l a t i o n s h i p . As h a s b e e n 70. shown, t h e s e t h r e e m o r p h o l o g i c a l f e a t u r e s a r e i n t e r - r e l a t e d i n c o n i f e r c a m b i a l m o r p h o l o g y w i t h l e n g t h f a c t o r a p p e a r i n g t o be t h e more d o m i n a n t one. One a d v a n t a g e o f t h i s a p p r o a c h i s t h a t , b a s e d on t h e l e n g t h f a c t o r a l o n e , t h e amount o f c a r b o h y d r a t e i n any t r a c h e i d c o u l d be e s t i m a t e d i r r e s p e c t i v e o f t h e o r i g i n o f wood. I n o t h e r words ,• f o r . p r a c t i c a l - c o n s i d e r a t i o n , t h e c a r - b o h y d r a t e . f r a c t i o n w o u l d be s i m i l a r f o r . t r a c h e i d s o f t h e same •length.;, i n t h e c o n i f e r o u s woods examined.. "• - . - . - - • I n v i e w o f s u c h a s t r o n g r e l a t i o n s h i p between c e l l l e n g t h and w e i g h t , and a p r i o r i c o n s i d e r a t i o n s f r o m c e l l m o r p h o l o g y , i t i s r e a s o n a b l e t o h y p o t h e s i z e t h a t t h e amount o f c a r b o h y d r a t e f r a c t i o n i n an i n d i v i d u a l t r a c h e i d i s a f u n c t i o n o f i t s l e n g t h . I t i s n o t known w h e t h e r t h e r e l a t i o n s h i p i s a d i r e c t one, i n t h a t t h e amount o f c a r b o h y d r a t e f r a c t i o n depends on t h e l e n g t h o f t h e c e l l ; o r w h e t h e r i t i s an i n d i r e c t one, i n t h a t b o t h v a r i a b l e s r e s p o n d t o some f i n i t e . f a c t o r . T h i s i s d i f f i c u l t t o a s s e s s , s i n c e f a c t o r s w h i c h i n f l u e n c e t h e e x t e n t o f c e l l e n l a r g e m e n t and w a l l d e v e l o p m e n t f o l l o w i n g c e l l d i v i s i o n a r e s t i l l n o t f u l l y u n d e r s t o o d . As p o i n t e d o u t by Bannan ( 8 ) , e v e n t u a l s i z e o f d e r i v e d c o n i f e r o u s e l e m e n t s i s d e t e r m i n e d l a r g e l y by s i z e o f t h e o r i g i - n a t i n g f u s i f o r m c a m b i a l c e l l s , p o s t - d i v i s i o n a l e l o n g a t i o n b e i n g r e l a t i v e l y s l i g h t i n c o n i f e r s . I n an e a r l i e r work Bannan and B a y l y (10) showed t h a t , i n c o n i f e r s , c e l l s i z e and e x t e n t o f r a y . c o n t a c t s a r e i m p o r t a n t f a c t o r s f o r s u r v i v a l o f c a m b i a l d a u g h t e r c e l l s . The l o n g e s t c e l l s w i t h t h e b e s t r a y c o n t a c t s have b e s t o p p o r t u n i t y t o s u r v i v e and d e v e l o p , whereas t h e s h o r t e s t f a i l o r f o r m new r a y i n i t i a l s . T h i s c r e a t e s an e f f e c - t i v e f e e d - b a c k r e g u l a t i v e s y s t e m w h i c h p r o v i d e s f o r an e q u i d i s - t a n t i n i t i a t i o n o f new r a y s ( 1 6 7 ) . The i m p o r t a n c e o f r a y s t o f u s i f o r m i n i t i a l s l i e s i n t h e i r f u n c t i o n o f t r a n s f e r and s t o r - age o f n u t r i e n t s , r e q u i r e d f o r t h e s u s t e n a n c e , g r o w t h , and d i v i s i o n o f t h e i n i t i a l s ; where t h e l o n g e s t i n i t i a l s a r e w e l l p r o v i d e d w i t h r a y c o n t a c t s t h e y p o s s i b l y h a v e a c c e s s t o abun- d a n t n u t r i e n t s . A p r o p e r b a l a n c e o f r a y c o n t a c t s commensurate w i t h t r a c h e i d l e n g t h , w o u l d p r o v i d e t h e d e r i v e d c e l l s w i t h s u f f i c i e n t n u t r i e n t s f o r t h e b u i l d - u p o f a r i g i d framework as t h e c e l l m a t u r e s . I t a p p e a r s f r o m t h e s e c o n s i d e r a t i o n t h a t a d i r e c t r e l a t i o n s h i p may e x i s t b etween t h e amount o f c a r b o h y d r a t e f r a c t i o n and t r a c h e i d l e n g t h . The r e s u l t s o b t a i n e d h e r e i n a r e t h u s c o n s i s t e n t w i t h f i n d i n g s r e p o r t e d by o t h e r s , i n t h a t t h e p h y s i o l o g i c a l mechanisms g o v e r n i n g t h e l e n g t h - w e i g h t r e l a t i o n - s h i p s a r e h i s t o r i e s o f g r o w t h r e l a t i o n s h i p s p o s s i b l y p r e d e t e r - m i n e d i n t h e o r i g i n a t i n g i n i t i a l s , a c c o r d i n g t o t h e i r s i z e . The r e l a t i o n s h i p d e s c r i b e d above has a n o t h e r i m p o r t a n t i m p l i c a t i o n i n t h a t l o n g e r t r a c h e i d s h a v e a g r e a t e r w e i g h t p e r u n i t l e n g t h t h a n s h o r t e r ones (see F i g u r e s 10 and 1 1 ) , a c o n c l u - s i o n a l s o r e a c h e d e a r l i e r by C l a r k ( 2 4 ) , who u s e d f i b r e ' c o a r s e - n e s s ' t o d e t e r m i n e . f i b r e w£'i';ght p e r u n i t l e n g t h o f c o m m e r c i a l s u l p h i t e p u l p s a m p l e s o f c o n i f e r o u s wood. As n o t e d , f o r l o n g e r t r a c h e i d s t h e a v e r a g e p e r c e n t a g e o f c e l l w a l l volume p e r u n i t l e n g t h w i l l be h i g h e r s i n c e l e n g t h i s c o r r e l a t e d p o s i t i v e l y w i t h d i a m e t e r (50, 54, 56, 158) and w a l l t h i c k n e s s (108, 1 4 0 ) . . A l s o , t h e p o s s i b i l i t y e x i s t s t h a t , f o r s h o r t e r t r a c h e i d s , t h e r e l a t i v e c o n t r i b u t i o n o f t i p s , w h i c h u n d o u b t e d l y , w e i g h l e s s , (146, p . 3 9 1 ) , i s more t h a n f o r l o n g e r t r a c h e i d s . I f t h i s i s s o , s u c h a r e l a t i o n s h i p may have i m p o r t a n t i m p l i - c a t i o n s f r o m t h e p r a c t i c a l p o i n t o f v i e w . A p p l i c a b i l i t y o f t h e R e l a t i o n s h i p A somewhat d i f f e r e n t a p p r o a c h was t a k e n t o a p p l y t h e r e l a t i o n s h i p as d e v e l o p e d t o p r o b l e m s s u c h as e f f e c t s o f f e r t i l i z e r s on wood p r o p e r t i e s . The r e l a t i o n s h i p has a l r e a d y b e e n a p p l i e d t o t e s t d i f f e r e n c e s between c o m p r e s s i o n wood t r a c h e i d s and t h o s e o f n o r m a l wood. I t i s t o be noted, however, t h a t c o m p r e s s i o n wood, on t h e one hand, i s a m o r p h o g e n e t i c phenomenon i n t r e e s f o r stem r e g u l a t o r y o r r e o r i e n t a t i o n f u n c - t i o n and, as s u c h , i t i s a n a t u r a l t h i n g t h a t t r e e s r e s p o n d a u t o m a t i c a l l y t o p r o d u c e s u c h wood. On t h e o t h e r hand, a f a c - t o r s u c h as c h e m i c a l f e r t i l i z a t i o n may be a r t i f i c i a l , c r e a t e d t o r a i s e t h e f e r t i l i t y l e v e l o f a s o i l i n o r d e r t o i n c r e a s e t h e y i e l d o r b r i n g n o n - p r o d u c t i v e l a n d i n t o p r o d u c t i o n . A l s o , f e r - t i l i z a t i o n a p p e a r s t o have l i t t l e d i r e c t e f f e c t on c a m b i a l a c t i v i t y o r wood d e v e l o p m e n t s i n c e t h e r e i s no f i r m e v i d e n c e t o i n d i c a t e t h a t t h e cambium c a n m o b i l i z e n u t r i e n t s d i r e c t l y 73. f r o m t h e t r a n s p i r a t i o n s t r e a m ( 7 0 ) . F e r t i l i z a t i o n e x e r t s i t s p r i m a r y o r d i r e c t e f f e c t on crown, and a l s o on r o o t d e v e l o p - • ment; and i t s e f f e c t on wood g r o w t h i s b o t h s e c o n d a r y t o and t h e r e s u l t o f crown r e s p o n s e s ( 7 9 ) . Thus i t a p p e a r s t h a t , f o l l o w i n g t h e a d d i t i o n o f a f e r t i l i z e r , a l l t h e p h y s i o l o g i c a l p r o c e s s e s i n t h e t r e e a r e a c t i v a t e d , so t o s p e a k . From t h e above v i e w p o i n t , i t w o u l d be o f i n t e r e s t t o see what happens t o t h e t r a c h e i d l e n g t h - w e i g h t r e l a t i o n s h i p t h a t i s o p e r a t i v e i n n o r m a l wood i n r e s p o n s e t o t h e a c t i o n o f f e r t i l i z a t i o n . T h r e e p o s s i b i l i t i e s e x i s t : " (a) T h e r e m i g h t be a g a i n i n c a r b o h y d r a t e amount p e r u n i t l e n g t h of t r a c h e i d o w i n g t o t h e i n c r e a s i n g a v a i l a b i l i t y o f p h o t o s y n t h a t e , s i n c e t h e r e w i l l be an i n c r e a s e i n p h o t o s y n t h e t i c s u r f a c e and p o s - s i b l y t h e p h o t o s y n t h e t i c e f f i c i e n c y o f f o l i a g e (18, 139); (b) d e c r e a s e i n c e l l w e i g h t p e r u n i t l e n g t h due t o o v e r p r o d u c t i o n o f c e l l s i n a g i v e n p e r i o d ; and (c) no c h ange i n s p i t e o f o v e r p r o d u c t i o n s i n c e t h e t r e e r e v e r t s t o a t e m p o r a r y p e r i o d o f j u v e n i l i t y by p r o d u c i n g s h o r t e r c e l l s c o n s e q u e n t t o i n c r e a s e i n t h e r a t e o f a n t i c l i n a l c a m b i a l c e l l d i v i s i o n s . . The b e s t way t o examine t h e s e p o s s i b i l i t i e s w o u l d be t o compare t h e t r a c h e i d l e n g t h - w e i g h t r e l a t i o n s h i p s o b t a i n e d f r o m t h e f e r t i l i z e d t r e e s , .with t h a t o b t a i n e d f o r t h e c ombined d a t a f o r c o n i f e r s . F o r o b v i o u s r e a s o n s , t h i s c a n n o t be done i n t h e p r e s e n t c a s e , i . e . , b e c a u s e o f ' s l o p e ' and ' l e v e l ' d i f f e r e n c e s b e t w e e n some o f t h e s p e c i e s s t u d i e d . I n v i e w o f t h i s , c o m p a r i - s o n s c a n o n l y be made w i t h t h e r e l a t i o n s h i p o b t a i n e d f o r n o r m a l D o u g l a s - f i r wood. B e f o r e i n v e s t i g a t i n g t h e s e a s p e c t s , i t w o u l d be a p p r o p - r i a t e t o t r a c e t h e c h a n g i n g p a t t e r n o f t r a c h e i d w e i g h t and l e n g t h , s t a r t i n g b e f o r e t h e f e r t i l i z a t i 6 n p e r i o d . S u c h a p a t t e r n i s shown i n F i g u r e 14, f o r a 2 9 - y e a r ^ o l d D o u g l a s - f i r t r e e w h i c h was f e r t i l i z e d a t a g e s 21 and 22. F o r c o m p a r i s o n , d a t a f o r i n c r e m e n t w i d t h , and mean t r a c h e i d l e n g t h b a s e d on 50 m a c e r a t e d t r a c h e i d s p e r i n c r e m e n t , w e r e a l s o p l o t t e d i n t h e same f i g u r e . I t c a n be s e e n t h a t t r a c h e i d w e i g h t o f b o t h h o l o - and a l p h a - c e l l u l o s e s k e l e t o n s d e c r e a s e d o r r e m a i n e d s t a t i c i m m e d i a t e l y f o l l o w i n g f e r t i l i z a t i o n . I n t h r e e o f t h e f o u r c a s e s t h i s d e c - r e a s e i n w e i g h t c o n t i n u e d f o r two y e a r s f o l l o w i n g t h e y e a r o f a p p l i c a t i o n ( o r one y e a r a f t e r r e f e r t i l i z a t i o n ) ; t h e r e a f t e r t h e r e was a s u d d e n i n c r e a s e i n w e i g h t f o l l o w e d a g a i n by a d e - c r e a s e due t o t h e f o r m a t i o n o f c o m p r e s s i o n wood. The o n l y e x c e p t i o n s w e r e t h e l a t e w o o d h o l o c e l l u l o s e s k e l e t o n s , w h i c h showed i n c r e a s e i n w e i g h t a f t e r a t e m p o r a r y d e c r e a s e , f o r o n l y one y e a r f o l l o w i n g t h e y e a r o f f e r t i l i z a t i o n . S u b s e q u e n t l y , t h e i r b e h a v i o u r was s i m i l a r t o t h e g e n e r a l p a t t e r n d e s c r i b e d a b o v e . T h i s i n c r e a s e i n l a t e w o o d t r a c h e i d w e i g h t ( y e a r 23) may h a v e b e e n due t o t h e i n c r e a s e d a v a i l a b i l i t y o f c u r r e n t p h o t o s y n t h a t e t o t h e c a m b i a l r e g i o n , a f t e r a l r e a d y m e e t i n g t h e new n e e d l e r e q u i r e m e n t s (49, 8 0 ) . F u r t h e r , t h i s was p r i m a r i l y an i n c r e a s e i n t h e h e m i c e l l u l o s e f r a c t i o n s i n c e no i n c r e a s e i s e v i d e n t f o r a l p h a - c e l l u l o s e s k e l e t o n w e i g h t s i n t h a t y e a r . B o t h mean t r a c h e i d l e n g t h f o r t h e i n c r e m e n t ( F i g u r e 14) and a v e r a g e t r a c h e i d l e n g t h o f a l l t h e t r a c h e i d s w e i g h e d ( T a b l e s I I I and IV) f o l l o w e d t h e same g e n e r a l p a t t e r n s n o t e d f o r t r a c h e i d w e i g h t . As e x p e c t e d , t h e r e was a s t e e p i n c r e a s e i n i n c r e m e n t w i d t h f o r t h e y e a r s 21 a n d 22, f o l l o w i n g f i r s t f e r t i l i z a t i o n ( y e a r 2 1 ) ; s u b s e q u e n t t o y e a r 23 t h e r e was no f u r t h e r i n c r e a s e , b u t g r o w t h r a t e was s t i l l q u i t e h i g h as c o m p a r e d t o t h a t b e f o r e f e r t i l i z a t i o n . The a b o v e p a t t e r n p r o v i d e s e v i d e n c e o f c h a n g e s i n t r a c h e i d w e i g h t f o l l o w i n g f e r t i l i z a t i o n . H o w e v e r , i t i s n o t c l e a r w h e t h e r t h e a p p a r e n t c h a n g e s a r e i n d e p e n d e n t o f v a r i a t i o n i n t r a c h e i d l e n g t h o r w h e t h e r t h e y a r e d i r e c t l y r e l a t e d t o t r a c h e i d l e n g t h . I n o r d e r t o e x a m i n e t h e v a r i a b i l i t y i n t h i s t r e e and o t h e r s , more f e r t i l i z e d wood m a t e r i a l o f D o u g l a s - f i r , a v a i l a b l e i n t h e f o r m o f i n c r e m e n t c o r e s f r o m a n o t h e r s t u d y ( 1 1 9 ) , was e m p l o y e d . As n o t e d , t h r e e f e r t i l i z e r c o m p o s i t i o n s h a d b e e n s e l e c t e d , n a m e l y U r e a , NPK„ a n d ( N H ^ ^ S C ^ . Two i n c r e - m e n t s , i . e . , 15 and 16 ( i n c l u d i n g a n d f o l l o w i n g r e f e r t i l i z a t i o n ) , f r o m one t r e e e a c h o f t h e s e t r e a t m e n t s w e r e s t u d i e d . The r e a s o n s f o r s e l e c t i o n o f t h e s e t r e a t m e n t s w e r e t h a t , w h i l e a l l g e n e r a t e d r e s p o n s e i n i n c r e m e n t w i d t h , t h e y showed a l s o d e f i n i t e i n c r e a s e ( U r e a ) o r d e c r e a s e (NPK, (NH.)„S0.) i n s p e c i f i c g r a v i t y f o l l o w i n g f e r t i l i z a t i o n t r e a t m e n t s . Some s e l e c t e d wood a n d t r a c h e i d p r o p e r t i e s f o r t h e t h r e e t r e e s a r e g i v e n i n T a b l e X I . The m a i n p u r p o s e h e r e was t o d e t e r m i n e t h e c a r b o h y d r a t e s k e l e - - t o n w e i g h t s p e r u n i t l e n g t h , a n d t o r e l a t e t h e s e t o c h a n g e s o b s e r v e d i n g r o s s wood s p e c i f i c g r a v i t y . I f j u a n d Kennedy (60) showed a h i g h c o r r e l a t i o n b e t w e e n s p e c i f i c g r a v i t y . a n d p e r c e n t a g e o f c e l l w a l l , w h i c h w o u l d s u g g e s t a p o s s i b l e c o n - v e r s i o n f r o m c e l l w a l l a r e a t o d e n s i t y v a l u e s . S i n c e c e l l l e n g t h i s c o r r e l a t e d w i t h c e l l d i a m e t e r a n d w a l l t h i c k n e s s as w e l l as w e i g h t , a p o s s i b i l i t y o f r e l a t i n g c e l l w e i g h t s t o s p e c i f i c g r a v i t y may be e x p l o r e d . S u c h a r e l a t i o n s h i p h a s a l r e a d y b e e n i n d i c a t e d f o r t h e 5 0 0 - y e a r - o l d D o u g l a s - f i r t r e e ( F i g u r e 1 2 ) . R e l a t i o n s h i p s b e t w e e n t r a c h e i d l e n g t h a n d w e i g h t , i n 2 t h e f e r t i l i z e d D o u g l a s - f i r t r e e s , a r e i n d i c a t e d i n F i g u r e s 15 t o 18, a l o n g w i t h t h a t o b t a i n e d f o r D o u g l a s - f i r n o r m a l wood. F o r e f f e c t i v e c o m p a r i s o n , t h e a n a l y s e s done i n e a c h c a s e a r e a l s o g i v e n i n e a c h f i g u r e . _A summary o f t h e o v e r a l l c o m p a r i s o n s among t h e t r e a t e d a n d u n t r e a t e d D o u g l a s - f i r t r e e s i s shown i n T a b l e X I I . I t c a n be i n f e r r e d f r o m t h e c o m b i n e d a n a l y s i s ( T a b l e X I I A) t h a t no s i g n i f i c a n t d i f f e r e n c e e x i s t s , f o r h o l o - c e l l u l o s e s k e l e t o n s , b e t w e e n t h e n o r m a l wood t r a c h e i d w e i g h t - 2 I t a p p e a r s f r o m F i g u r e 15 t h a t t h e r e g r e s s i o n l i n e o f h o l o c e l l u l o s e t r a c h e i d s k e l e t o n s o f t h e U r e a t r e a t e d t r e e i s d i f f e r e n t f r o m t h a t o f n o r m a l D o u g l a s - f i r . S t a t i s t i c a l a n a l y s e s , h o w e v e r , i n d i c a t e t h a t t h e a p p a r e n t d i f f e r e n c e s a r e n o t s i g n i f i - c a n t . T h i s i s t r u e b e c a u s e t h e s t a n d a r d e r r o r o f e s t i m a t e (SEE) o f n o r m a l D o u g l a s - f i r ( F i g u r e 9 — H o l o c e l l u l o s e ) i s a b o u t t h r e e t i m e s t h a t o f t h e U r e a t r e a t e d t r e e a n d t h u s i t w o u l d i n c l u d e a l l v a l u e s o f t h e l a t t e r . l e n g t h r e l a t i o n s h i p a n d t h a t o f f e r t i l i z e d wood. When e a c h t r e a t m e n t was c o m p a r e d i n d i v i d u a l l y w i t h n o r m a l D o u g l a s - f i r ( s e e F i g u r e s 15 t o 18; H o l o c e l l u l o s e ) , w i t h one e x c e p t i o n ( N H 4 N 0 3 t r e a t m e n t — F i g u r e 1 8 ) , no s i g n i f i c a n t d i f f e r e n c e c o u l d be o b s e r v e d . C o n t r a r y t o t h e a b o v e , t h e c o m b i n e d a n a l y s i s f o r a l p h a - c e l l u l o s e s k e l e t o n s p o i n t s o u t s i g n i f i c a n t d i f f e r e n c e s ( T a b l e X I I A) among and w i t h i n t h e f i v e D o u g l a s - f i r t r e e s . The same b e h a v i o u r was n o t e d when t r e a t m e n t s w e r e c o m p a r e d i n d i v i - d u a l l y w i t h n o r m a l D o u g l a s - f i r ( F i g u r e s 16 t o 1 8 ) . I n t e r e s t i n g l y , a l p h a - c e l l u l o s e s k e l e t o n s ' f r o m t r e a t m e n t U r e a d i d n o t d i f f e r e i t h e r i n s l o p e o r l e v e l f r o m n o r m a l D o u g l a s - f i r ( F i g u r e 1 5 ) . C o m p a r i s o n s b e t w e e n t h e t r e a t m e n t s ( T a b l e X I I B ) , f o r h o l o c e l l u l o s e s k e l e t o n s , i n d i c a t e t h a t t h e r e i s no s i g n i f i c a n t d i f f e r e n c e i n s l o p e s , t h a t i s , t h e y a r e p a r a l l e l . T h i s i m p l i e s t h a t t h e b e h a v i o u r ( r e s p o n s e ) o f t h e d i f f e r e n t t r e a t m e n t s , w i t h r e s p e c t t o t r a c h e i d l e n g t h - w e i g h t r e l a t i o n s h i p , i s s i m i l a r . B u t t h e m a g n i t u d e o f r e s p o n s e i n t h e s e t r e a t m e n t s i s d i f f e r e n t s i n c e t h e y d i f f e r e d i n l e v e l s . The o n l y e x c e p t i o n was b e t w e e n U r e a and NPK w h i c h d i d n o t d i f f e r i n e i t h e r l e v e l s o r s l o p e s . I n v i e w o f t h e l e v e l d i f f e r e n c e s , i t c a n be c o n c l u d e d t h a t t h e s e t r e a t m e n t s may d i f f e r i n t h e i r e f f e c t s o n wood p r o p e r t i e s . T h u s , i f t h e r e g r e s s i o n e q u a t i o n s o f b e s t f i t ( F i g u r e s 15 t o 18) a r e e x a m i n e d f o r p r e d i c t e d v a l u e s , U r e a t r e a t m e n t shows h i g h e r v a l u e s i n t h a t h o l o c e l l u l o s e t r a c h e i d s a r e h e a v i e r p e r u n i t l e n g t h , t h a n c o m p a r a b l e o n e s f r o m t h e o t h e r t r e a t m e n t s . I t i s 78. t h e r e f o r e p o s s i b l e t h a t changes a s s o c i a t e d w i t h c e l l w e i g h t a r e r e s p o n s i b l e , a l o n g w i t h o t h e r f a c t o r s ( s e e T a b l e X I ) , f o r t h e o b s e r v e d changes i n g r o s s wood s p e c i f i c g r a v i t y i n t h e s e f e r t i l i z e d t r e e s . The b e h a v i o u r o f a l p h a - c e l l u l o s e s k e l e t o n s f r o m d i f - f e r e n t t r e a t m e n t s i s somewhat d i f f e r e n t t h a n t h a t o f h o l o c e l l u - l o s e s k e l e t o n s . S i g n i f i c a n t d i f f e r e n c e s e x i s t between t h e r e g r e s s i o n l i n e s o f t r e a t m e n t s U r e a , NPK and (NH^) i n t h e i r r e s p e c t i v e s l o p e s , w h i l e U r e a and NH^NO^ d i d n o t show s u c h d i f f e r e n c e ( T a b l e X I I B ) . W i t h r e g a r d t o l e v e l s , U r e a d i f f e r e d s i g n i f i c a n t l y f r o m (NH^) 2S(-)4 k u t n o t f r o m NPK and NH^NO-j. H e r e a g a i n , when c o m p a r i s o n s a r e made on t h e b a s i s o f r e g r e s s i o n e q u a t i o n s o f b e s t f i t ( F i g u r e s 15 t o 1 8 ) , t h e U r e a t r e a t e d t r e e showed h i g h e r v a l u e s ( p r e d i c t e d t r a c h e i d w e i g h t ) , p e r u n i t l e n g t h , t h a n t h o s e o f t h e o t h e r t r e a t m e n t s . I n t h i s r e s p e c t , r e s u l t s f o r a l p h a - c e l l u l o s e s k e l e t o n s a r e s i m i l a r t o t h e f i n d i n g s r e p o r t e d f o r h o l o c e l l u l o s e , t h u s c o n f i r m i n g t h e argument a d v a n c e d f o r t h e r e l a t i o n s h i p between i n d i v i d u a l t r a c h e i d w e i g h t and g r o s s , wood s p e c i f i c g r a v i t y . A l t h o u g h t h e above comments r e f e r t o t r a c h e i d w e i g h t - l e n g t h r e l a t i o n s h i p s i n f e r t i l i z e d wood, t h e y a l s o p r o v i d e a d d i - t i o n a l p r o o f f o r t h e t h e s i s h y p o t h e s i s i n t h a t t h e p h y s i o l o g i c a l mechanism g o v e r n i n g t h e t r a c h e i d w e i g h t - l e n g t h r e l a t i o n s h i p s i n c o n i f e r s i s o n c e a g a i n t h e same i n t h e f e r t i l i z e d t r e e s . From t h e b i o l o g i c a l v i e w p o i n t , t h i s means t h a t w h i l e r e a d j u s t m e n t s f o r i n c r e a s e d g r o w t h r a t e a r e t a k i n g p l a c e , t h e s u g g e s t e d r e l a t i o n s h i p i s m a i n t a i n e d . O b v i o u s l y t h i s i s a c h i e v e d by a t e m p o r a r y r e v e r s a l t o j u v e n i l i t y , s o t h a t c h a n g e s i n t r a c h e i d w e i g h t a r e b a s i c a l l y a r e f l e c t i o n o f c h a n g e s i n t r a c h e i d l e n g t h . W h i l e t h i s seems t o be t h e g e n e r a l c a s e , w i t h i n t h e l i m i t s o f t h e s u g g e s t e d r e l a t i o n s h i p t h e r e w e r e some s t a t i s t i c a l d i f f e r - e n c e s among t h e t r e a t e d t r e e s i n t h e i r m a g n i t u d e o f r e s p o n s e ( w e i g h t p e r u n i t l e n g t h o f t r a c h e i d ) i n d i c a t i n g t h e p o s s i b i l i t y f o r d i f f e r e n t i a l e f f e c t s o f f e r t i l i z e r s o n wood p r o p e r t i e s . Many s t u d i e s on t h e e f f e c t s o f f e r t i l i z a t i o n on wood p r o p e r t i e s h a v e shown a d e c r e a s e i n t r a c h e i d l e n g t h a s s o c i a t e d w i t h i n c r e a s e d g r o w t h r a t e ( 7 1 , 108, 1 1 7 ) . I t i s e v i d e n t t h a t a t r e e h a s t h i s r e m a r k a b l e t y p e o f m e c h a n i s m o f r e v e r t i n g t o t e m p o r a r y j u v e n i l i t y as a r e s u l t o f a n i n c r e a s e d r a t e o f a n t i - c l i n a l d i v i s i o n s i n t h e c a m b i a l c e l l s . T h e s e a s p e c t s a n d r e - l a t e d p r o c e s s e s a r e w e l l d o c u m e n t e d i n : t h e e x t e n s i v e w o r k s o f B a n n a n ( 8 ) . I n v i e w o f s u c h e f f e c t s , c o m p a r i s o n s w e r e made b e t w e e n b e h a v i o u r o f f e r t i l i z e d wood and t h a t o f j u v e n i l e wood ( T a b l e X I I B ) . From t h e s e a n a l y s e s , i t i s c l e a r t h a t no s i g n i - f i c a n t d i f f e r e n c e s c a n be f o u n d e i t h e r i n s l o p e s o r l e v e l s o f h o l o c e l l u l o s e r e g r e s s i o n l i n e s , b e t w e e n " j u v e n i l e wood t r a c h e i d s and t h o s e f r o m t h e t r e a t m e n t s U r e a , NPK and (NH^^SO^. B u t a s i g n i f i c a n t d i f f e r e n c e e x i s t s ( i n s l o p e s o n l y ) when c o m p a r i n g j u v e n i l e wood t r a c h e i d s w i t h t h o s e f r o m t r e a t m e n t NH^NO^ . T h i s d i f f e r e n c e i n b e h a v i o u r c o u l d be a t t r i b u t e d t o t h e age f a c t o r o f t h i s t r e e . F o r e x a m p l e , t h e NH^NO^ f e r t i l i z e d t r e e was 21 y e a r s o l d a t t h e t i m e o f f i r s t f e r t i l i z a t i o n ( F i g u r e 1 4 ) , 80. w h e r e a s t h e U r e a , NPK, (NH^) t r e e s w e r e age 13 ( v e r y c l o s e t o j u v e n i l e p e r i o d — s e e T a b l e X I ) when f e r t i l i z e d . C o n s e q u e n t l y h o l o c e l l u l o s e t r a c h e i d s k e l e t o n s f r o m t r e a t m e n t NH^NO^ d i f f e r e d f r o m t h o s e o f j u v e n i l e wood. A n a l y s e s f o r a l p h a - c e l l u l o s e i n d i c a t e no s i g n i f i c a n t d i f f e r e n c e s i n r e g r e s s i o n l i n e s l o p e s b e t w e e n t h e j u v e n i l e wood t r a c h e i d s and t h o s e f o r t h e t r e a t m e n t s U r e a , NPK, and ( N H ^ ) 2 S 0 4 ( T a b l e X I I B ) . Once a g a i n r e s u l t s f o r t r e a t m e n t NH^NO^ d i f f e r e d s i g n i f i c a n t l y i n s l o p e f r o m t h o s e o f j u v e n i l e wood, s u g g e s t i n g t h a t t h e s e two t r a c h e i d p o p u l a t i o n s d i f f e r . The i n t e r e s t i n g p o i n t , h o w e v e r , i s t h e s i g n i f i c a n t d i f f e r e n c e i n l e v e l s o f r e g r e s s i o n l i n e s b e t w e e n j u v e n i l e wood a nd t r e a t m e n t s U r e a a n d NPK. I n c o n t r a s t t o t h i s , a l p h a - c e l l u l o s e s k e l e t o n s f r o m t r e a t m e n t (NH^) 2^4 n o t show s i g n i f i c a n t d i f f e r e n c e f r o m t h o s e o f j u v e n i l e wood e i t h e r i n s l o p e s o r l e v e l s o f r e g r e s s i o n l i n e s . The i m p l i c a t i o n s o f t h e s e f i n d i n g s a r e i m p o r t a n t i n t h a t t r e a t m e n t U r e a and NPK d i f f e r e d f r o m (NH^)2 S 04 w i t h r e s p e c t t o r e s u l t i n g c a r b o h y d r a t e f r a c t i o n p e r u n i t l e n g t h o f t r a c h e i d . F u r t h e r m o r e , when c o m p a r e d w i t h n o r m a l D o u g l a s - f i r , o n l y U r e a ( F i g u r e 15) d i d n o t show any s i g n i f i c a n t d i f f e r e n c e , w h e r e a s b o t h NPK and ( N H ^ ) 2 S 0 4 ( F i g u r e s 16 a n d 17) d i f f e r e d s i g n i f i c a n t l y i n t h e amount o f a l p h a - c e l l u l o s e p e r u n i t l e n g t h o f t r a c h e i d . The b e h a v i o u r o f t r a c h e i d s k e l e t o n s f r o m t r e a t m e n t NH^NO^ i s somewhat d i f f e r e n t f r o m t h e o t h e r s . When c o m p a r e d t o n o r m a l D o u g l a s - f i r , t h e NH.NO-. t r e a t e d t r e e ( F i g u r e 18) showed 81. s i g n i f i c a n t d i f f e r e n c e s i n s l o p e s f o r b o t h h o l o - a n d a l p h a - c e l l u l o s e s k e l e t o n s , s u g g e s t i n g t h a t t h e t r a c h e i d p o p u l a t i o n s f r o m t h e s e two t r e e s d i f f e r . I n t e r e s t i n g l y , t h e u p p e r p a r t o f t h e c u r v e i n F i g u r e 18 ( H o l o c e l l u l o s e — w h e r e X = 3.92) i n d i - . c a t e s t h a t t h e s e t r a c h e i d s may h a v e more h o l o c e l l u l o s e p e r u n i t l e n g t h t h a n c o m p a r a b l e n o r m a l wood t r a c h e i d s . S u c h b e h a v i o u r may be a t t r i b u t e d t o t h e i n c r e a s e i n w e i g h t n o t e d i n t h e l a t e w o o d h o l o c e l l u l o s e s k e l e t o n s ( y e a r s 23 a n d 2 4 - — F i g u r e 1 4 ) , f o l l o w i n g r e f e r t i l i z a t i o n . I n c o n c l u s i o n , i t c a n be s a i d t h a t t h e b a s i c r e l a t i o n - s h i p s u g g e s t e d f o r n o r m a l wood, b e t w e e n l e n g t h a n d w e i g h t , i s m a i n t a i n e d i n t h e wood f r o m f e r t i l i z e d t r e e s ; s u c h t r e e s a d j u s t q u i c k l y t o t h e i n c r e a s e d g r o w t h r a t e by a t e m p o r a r y r e v e r s a l t o j u v e n i l i t y and p r o d u c e s h o r t e r t r a c h e i d s . I n a d d i t i o n , t h e p o s s i b i l i t y e x i s t s f o r a d e c r e a s e i n w e i g h t ( a l p h a - c e l l u l o s e ) p e r u n i t l e n g t h as e x e m p l i f i e d by t r e a t m e n t s NPK and (NH^^SO^. The p e r i o d o f t e m p o r a r y j u v e n i l i t y and t h e d e g r e e o f r e d u c t i o n i n t r a c h e i d l e n g t h , may w e l l be r e l a t e d t o r a t e o f i n c r e a s e i n g r o w t h as w e l l as t o o t h e r f a c t o r s s u c h as i n i t i a l t r e e v i g o r and a g e . Where t h e t r e e s a r e g r o w i n g v i g o r o u s l y a n d s t i l l i n t h e p e r i o d o f r a p i d g r o w t h ( 4 6 a ) , t h e y m i g h t r e c o v e r s p e e d i l y f r o m t h i s t e m p o r a r y r e v e r s a l t o j u v e n i l i t y . I f t h e t r e e s a r e o l d and p a s t t h i s p e r i o d , i t may t a k e l o n g e r b e f o r e t h e y r e a c h t h e e x p e c t e d norm i n t r a c h e i d l e n g t h - w e i g h t . I n t h e p r e s e n t i n s t a n c e t h e f e r t i l i z e d t r e e s u s e d w e r e y o u n g , a t t h e t i m e o f f e r t i l i z a t i o n , and i t i s n o t known w h e t h e r t h e r e p o r t e d t r e n d s 82. i n t r a c h e i d w e i g h t w i l l be t h e same f o r o l d e r s t a n d s . N e v e r - t h e l e s s , t h e r e s u l t s o b t a i n e d do n o t i n d i c a t e any d r a s t i c c h a n g e i n f i b r e p r o p e r t i e s e v e n i n o l d e r t r e e s , when f e r t i l i z e r i s a p p l i e d i n m o d e r a t i o n and w i t h due c o n s i d e r a t i o n t o o t h e r s i l v i c u l t u r a l and management p r a c t i c e s ( 7 9 ) . Some P r a c t i c a l C o n s i d e r a t i o n s Two p r a c t i c a l a s p e c t s w i l l be d i s c u s s e d i n t h i s s e c - t i o n , n a m e l y , e x p e r i m e n t a l y i e l d o f c a r b o h y d r a t e f r a c t i o n a nd t h e i m p l i c a t i o n o f t h e t r a c h e i d l e n g t h - w e i g h t r e l a t i o n s h i p o b t a i n e d , t o a r e a s o f wood and p u l p s c i e n c e and i n d u s t r y . Y i e l d o f h o l o c e l l u l o s e f r a c t i o n o b t a i n e d by t h e p r e s e n t m e t h o d , u s i n g p e r a c e t i c a c i d (and N a B H 4 r e d u c t i o n ) , h a s b e e n r e p o r t e d by t h e o r i g i n a t o r (83) t o be somewhat i n t e r m e d i a t e b e t w e e n c h l o r i t e h o l o c e l l u l o s e (163) and t h a t o f T A P P I ( c h l o r i n e - e t h a n o l a m i n e (128)) m e t h o d s . However, t h e p e r a c e t i c a c i d m e t h o d a d o p t e d h a s b e e n s u g g e s t e d t o g i v e a s u p e r i o r h o l o c e l l u l o s e w i t h a D.P. o f 3050 t o 3600, 93 t o 100 p e r c e n t r e c o v e r y o f m a j o r p o l y s a c c h a r i d e s and minimum o f c a r b o x y l g r o u p s ( 8 3 ) . The y i e l d s o b t a i n e d i n t h e p r e s e n t s t u d y f o r b o t h h o l o - and a l p h a - c e l l u l o s e ( A p p e n d i x I I ) , a r e c o m p a r a b l e t o t h o s e r e p o r t e d by o t h e r s (20, 123, 1 3 5 ) . T h e r e a p p e a r s t o b e some p e c u l i a r i t y f o r t h i s m e t hod ( p e r a c e t i c a c i d ) i n t h a t t h e h o l o c e l l u l o s e y i e l d s f o r e a r l y w o o d a r e s l i g h t l y h i g h e r t h a n t h o s e f r o m l a t e w o o d . S i m i l a r d i f f e r - e n c e s i n y i e l d h a v e a l s o b e e n " r e c o r d e d by L e o p o l d f o r l o b l o l l y 83. p i n e wood (83) . The r e a s o n f o r t h i s d i f f e r e n c e i s o b v i o u s s i n c e t h e c o o k i n g c y c l e s n e e d ed f o r e a r l y w o o d were o n l y t h r e e as compared t o f i v e f o r l a t e w o o d . B u t t h i s b e h a v i o u r does n o t seem t o i n f l u e n c e t h e a l p h a - c e l l u l o s e y i e l d s , w h i c h showed t h e e x p e c t e d d i f f e r e n c e i n t h a t l a t e w o o d v a l u e s were h i g h e r (Appen- d i x I I ) . C o r r e c t i o n s t o t h e w e i g h t o f i n d i v i d u a l t r a c h e i d s , b a s e d on y i e l d , h a v e n o t b e e n made s i n c e i t i s d i f f i c u l t t o a s s e s s s u c h m i n o r d i f f e r e n c e s i n s i n g l e t r a c h e i d w e i g h t s . I n a d d i t i o n , i t i s i n c o r r e c t t o assume t h a t d i f f e r e n t woods w i l l h a v e t h e same y i e l d , due t o i n h e r e n t v a r i a b i l i t i e s . As s u c h , c o m p a r i s o n s b a s e d on t h e a c t u a l v a l u e s o b t a i n e d w i l l be more m e a n i n g f u l and r e a l i s t i c t h a n t h o s e b a s e d on a d j u s t e d v a l u e s . However, i n t h i s work, c a r e was t a k e n t o s e e t h a t t h e same d e l i g n i f i c a t i o n c o n d i t i o n s were employed f o r a l l t h e woods S p o t c h e c k s o f some o f t h e p u l p samples i n d i c a t e d c o m p a r a b l e amounts o f l i g n i n ( A p p e n d i x I I ) . I n t h e l i g h t o f t h e p r e s e n t s t u d y , . i t a p p e a r s t h a t t h e l e n g t h - w e i g h t r e l a t i o n s h i p o f t h e c a r b o h y d r a t e s k e l e t o n s o b t a i n e d c a n be u s e d p r o f i t a b l y , by r e - e m p h a s i z i n g t h e i m p o r - . t a n c e o f t r a c h e i d l e n g t h f o r s e l e c t i o n o f i n d i v i d u a l s i n b r e e d i and v e g e t a t i v e p r o p a g a t i o n work. I t s i m p o r t a n c e t o t h e p u l p and p a p e r i n d u s t r y l i e s i n t h e j u d i c i o u s s e l e c t i o n o f i n d i - v i d u a l s w i t h i n s p e c i e s h a v i n g l o n g e r f i b r e s , n o t o n l y b e c a u s e o f t h e r e l a t i o n s h i p b etween f i b r e m o r p h o l o g y and p a p e r p r o p e r t i e s 84. (13, 29, 34, 1 5 0 ) , b u t a l s o b e c a u s e o f p o s s i b l e d i f f e r e n c e s i n c a r b o h y d r a t e y i e l d . I n a d d i t i o n , t h e r e l a t i o n s h i p between l e n g t h and w e i g h t i s a c u r v i l i n e a r one, meaning t h a t f o r e v e r y i n c r e a s e i n u n i t l e n g t h o f f i b r e , t h e r e w i l l be a d i s p r o p o r - t i o n a t e i n c r e a s e i n t h e amount o f c e l l u l o s e ( F i g u r e s 10, 1 1 ) . I t t h e r e f o r e a p p e a r s t h a t l o n g e r f i b r e s c o u l d r e s u l t i n a h i g h e r p u l p y i e l d due t o f e w e r end w a l l c o n t a c t a r e a s p e r g i v e n w e i g h t o r volume o f wood, w h i c h c o n s e q u e n t l y w o u l d r e s u l t i n l e s s l i g n i n (39) . T h i s b r i n g s up an i m p o r t a n t p o i n t r e l a t e d t o p u l p e v a l u a t i o n b a s e d on t h e number o f f i b r e s p e r gram, and i t s s i g n i f i c a n c e f o r a n a l y z i n g t h e ( p u l p ) c h a r a c t e r i s t i c s as a f f e c t e d by s p e c i e s , y i e l d o r p u l p i n g p r o c e s s . The i m p l i c a - t i o n s o f t h i s and r e l a t e d a s p e c t s h a v e been d i s c u s s e d r e c e n t l y by H o r n and Coens ( 5 8 ) . A t c o n s i d e r a b l e r i s k o f o v e r s i m p l i f i c a t i o n , i t c a n be s t a t e d t h a t i f l o n g e r f i b r e s r e s u l t i n more c e l l u l o s e c o n t e n t , p e r g i v e n w e i g h t o r volume o f wood, i t w o u l d seem a d v a n t a g e o u s t o grow t r e e s w i t h l o n g e r f i b r e s r a t h e r t h a n p r o p a g a t i n g s p e c i e s o r t r e e s w i t h s h o r t e r f i b r e s , i f t h e wood i s t o be u s e d i n t h e p u l p and p a p e r i n d u s t r y ; b r i e f l y , t h i s i s a h i g h y i e l d f o r e s t r y c o n c e p t . S i m i l a r l y , L e s l i e (84) s u g g e s t e d some y e a r s ago t h e c o n c e p t o f ' c e l l u l o s e f o r e s t r y , ' whereby f o r e s t s w o u l d be s c i e n t i f i c a l l y managed f o r t h e c o n t i n u o u s p r o d u c t i o n o f wood p o s s e s s i n g optimum p r o p e r t i e s f o r p a p e r a n d / o r p u l p m a k i n g . B r e e d i n g f o r f i b r e q u a l i t y i s j u s t i f i e d on t h e s e g r o u n d s . 85. M o r e o v e r , c o n s i d e r a b l e e v i d e n c e e x i s t s as t o t h e h e r i t a b l e n a t u r e o f f i b r e l e n g t h (8, 9, 37, 47, 61, 100, 154, 169, 1 7 0 ) . A l t h o u g h e s t i m a t e s f o r g r o s s wood c e l l u l o s e c o n t e n t do n o t show any s i g n i f i c a n t l y h e r i t a b l e n a t u r e , h i g h e r y i e l d s o f c e l l u l o s e c a n s t i l l be o b t a i n e d by b r e e d i n g f o r f i b r e l e n g t h s i n c e t h e i n d i v i d u a l f i b r e c o m p o s i t i o n i s h i g h l y s i g n i f i c a n t l y c o r r e l a t e d w i t h l e n g t h , as b r o u g h t o u t i n t h i s s t u d y . A n o t h e r p o i n t t h a t may m e r i t f u r t h e r c o n s i d e r a t i o n , i s s h o r t e n i n g r o t a t i o n age o f t h e s t a n d b e c a u s e o f i n c r e a s i n g p r e s s u r e f o r l a n d t o p r o d u c e maximum f i b r e r e t u r n s . The l e n g t h - w e i g h t r e l a t i o n s h i p o f t r a c h e i d s c a n be employed p r o - f i t a b l y , t o s u i t i n d i v i d u a l company r e q u i r e m e n t s , i n d e t e r - m i n i n g t h e r o t a t i o n age. T h i s p o i n t may be w o r t h i n v e s t i g a - t i o n i n p a r t i c u l a r f o r t h o s e c o u n t r i e s s e t t i n g up p l a n t a t i o n s o r c o u n t r i e s s t r i v i n g t o p r o d u c e enough wood t o meet t h e i n c r e a s i n g demands o f t h e p u l p and p a p e r i n d u s t r y . CHAPTER V RECOMMENDATIONS FOR FURTHER RESEARCH Much .of t h e i n f o r m a t i o n i n t h i s p a p e r on t r a c h e i d w e i g h t ( c a r b o h y d r a t e s k e l e t o n ) and l e n g t h r e l a t i o n s h i p s , r e p r e s e n t s a r e c o n n a i s s a n c e o f an o v e r a l l r e l a t i o n s h i p t h a t e x i s t s between and w i t h i n c o n i f e r o u s (stem) woods. I t s p r i m a r y o b j e c t i v e was t o d e t e r m i n e t h e more s t r i k i n g phenomenon e n c o u n - t e r e d , and p r o v i d e some b a s i s f o r s u b s e q u e n t and more d e t a i l e d i n v e s t i g a t i o n s . A l t h o u g h t h e p r o j e c t has b e e n d e s i g n e d t o i n c l u d e a w i d e r a n g e o f m a t e r i a l so t h a t c o n c l u s i o n s w i l l be a p p l i c a b l e t o c o n i f e r s i n g e n e r a l , many q u e s t i o n s n e e d i n g f u r t h e r s t u d y h a v e been r a i s e d . F o r example, a s t u d y o f t h e r e l a t i o n s h i p between i n d i v i d u a l f i b r e w e i g h t and i n t r a - i n c r e - m e n t a l s p e c i f i c g r a v i t y . m e r i t s f u r t h e r c o n s i d e r a t i o n . S i n c e a n a t o m i c a l e v i d e n c e i n d i c a t e s no s i g n i f i c a n t d i f f e r e n c e i n t h e a c t u a l amounts o f c e l l w a l l m a t e r i a l p e r t r a c h e i d b etween e a r l y w o o d and l a t e w o o d t r a c h e i d s , i t w o u l d be o f i n t e r e s t t o p u r s u e t h i s a s p e c t f u r t h e r w i t h r e g a r d t o t r a c h e i d w e i g h t s . A p r e l i m i n a r y a p p r o a c h has a l r e a d y b e e n made h e r e i n , b u t r e s u l t s a r e i n c o n c l u s i v e . O t h e r s t u d i e s t h a t n e e d f u r t h e r r e s e a r c h i n c l u d e t h e r e l a t i o n s h i p t h a t may e x i s t between i n d i v i d u a l f i b r e w e i g h t s and l e n g t h , f o r a n g i o s p e r m s , so as t o e v a l u a t e r e s u l t s f o r b o t h p r a c t i c a l and b i o l o g i c a l s i g n i f i c a n c e . 87. I t has b e e n o b s e r v e d i n t h i s s t u d y t h a t a r e d u c t i o n i n t r a c h e i d l e n g t h o c c u r s i n a l p h a - c e l l u l o s e t r a c h e i d s k e l e t o n s s u b s e q u e n t t o r e m o v a l o f t h e h e m i c e l l u l o s e f r a c t i o n i n t h e h o l o - c e l l u l o s e t r a c h e i d s k e l e t o n s . A s t u d y c o u l d be c o n d u c t e d t o t e s t w h e t h e r t h e r a t e ' o r d e g r e e o f r e d u c t i o n i s r e l a t e d t o t i s s u e t y p e ( e a r l y w o o d / l a t e w o o d ) and s p e c i e s . T h i s w o u l d p r o v i d e a more e x a c t l e n g t h - w e i g h t r e l a t i o n s h i p f o r a l p h a - c e l l u l o s e , so t h a t more r e a l i s t i c c o m p a r i s o n s c a n be made'with t h o s e o f h o l o c e l l u l o s e s k e l e t o n s . • T h i s i n f o r m a t i o n c o u l d be u s e f u l i n d e v e l o p m e n t a l s t u d i e s i n v o l v i n g t h e c h e m i c a l c h a n g es i n c e l l s , a c c o m p a n y i n g d i f f e r e n t i a t i o n i n t o x y l e m . F i n a l l y , - w e i g h ing«o.f:other c e l l e l e m e n t s s u c h as v e s s e l s and parenchyma c o u l d be done. S a s t r y and W e l l w o o d (118) s u g g e s - t e d s e v e r a l a r e a s o f r e s e a r c h where t r a c h e i d w e i g h t s c o u l d be employed p r o f i t a b l y . F o r example, s i n g l e f i b r e w e i g h t s c a n be u s e d i n d e t e r m i n i n g s t r e n g t h - w e i g h t o r s t r e n g t h / d e n s i t y r a t i o s f o r i n d i v i d u a l f i b r e s , d e n s i t y o f c e l l w a l l s i n c e l l w a l l p o r o s i t y s t u d i e s , i n q u a n t i t a t i v e s t u d i e s i n v o l v i n g wood f o r m a - t i o n , and a s s e s s i n g t h e e f f e c t s o f s i l v i c a l and e n v i r o n m e n t a l v a r i a b l e s on wood p r o p e r t i e s . The u n i q u e n e s s o f t h i s a p p r o a c h t h u s f a c i l i t a t e s d i r e c t c o m p a r i s o n o f c h a r a c t e r i s t i c s s u c h as s t r e n g t h p r o p e r t i e s , d i m e n s i o n s and c a r b o h y d r a t e c o n t e n t on t h e same f i b r e . CHAPTER VI CONCLUSION 1. A q u a r t z u l t r a - m i c r o b a l a n c e has b e e n d e v i s e d f o r a c c u r a t e w e i g h i n g o f s i n g l e c e l l s . The b a l a n c e has a w e i g h i n g r a n g e o f 0.06 t o 14 ug and a p r e c i s i o n ±0.03 y g . T h i s b a l a n c e was u s e d f o r q u a n t i t a t i v e d e t e r m i n a t i o n o f b o t h h o l o - and a l p h a - c e l l u l o s e i n i n d i v i d u a l t r a c h e i d s k e l e t o n s o f n i n e c o n i - f e r o u s woods, b e l o n g i n g t o n i n e d i f f e r e n t g e n e r a and s e v e n f a m i l i e s . 2. L e n g t h o f a t r a c h e i d and t h e amount.of c a r b o h y - d r a t e i n i t a r e h i g h l y s i g n i f i c a n t l y c o r r e l a t e d . The e s t i m a t e d v a r i a t i o n s a c c o u n t e d f o r i n h o l o c e l l u l o s e f r a c t i o n and t h a t o f a l p h a - c e l l u l o s e , by t h e l e n g t h f a c t o r a l o n e , were r e s p e c t i v e l y 91.9 and 95.7 p e r c e n t f o r a l l s p e c i e s s t u d i e d . A d i r e c t , p h y s i o l o g i c a l r e l a t i o n s h i p between l e n g t h o f t r a c h e i d and i t s w e i g h t ( c a r b o h y d r a t e s k e l e t o n ) i s s u g g e s t e d . 3. Between s p e c i e s d i f f e r e n c e s were o b s e r v e d w i t h r e s p e c t t o s l o p e s and l e v e l s o f r e g r e s s i o n l i n e s . D i f f e r e n c e s i n c e l l d i a m e t e r and w a l l t h i c k n e s s may have c o n t r i b u t e d t o t h i s s t a t i s t i c a l b e h a v i o u r . F o r p r a c t i c a l c o n s i d e r a t i o n s , t h e c o n i f e r o u s t r a c h e i d s c o u l d be g r o u p e d as one p o p u l a t i o n . S m a l l 2 s t a n d a r d d e v i a t i o n , - and h i g h l y s i g n i f i c a n t r v a l u e s , between t r a c h e i d w e i g h t and l e n g t h , s u p p o r t t h i s argument. 89. 4. The r a d i a l v a r i a t i o n o f s i n g l e t r a c h e i d w e i g h t f o l l o w e d an i n c r e a s i n g t r e n d up t o a b o u t 150 y e a r s and d e c r e a s e d s u b s e q u e n t l y i n t h e o v e r m a t u r e wood. 5. F o r t h e same t r a c h e i d l e n g t h , i n d i v i d u a l t r a c h e i d s o f j u v e n i l e wood h a d s i g n i f i c a n t l y l o w e r amounts o f a l p h a - c e l l u l o s e t h a n t h o s e f r o m m a t u r e and o v e r m a t u r e wood, whereas d i f f e r e n c e s were n o n s i g n i f i c a n t f o r h o l o c e l l u l o s e . 6. O v e r m a t u r e wood t r a c h e i d s had s i g n i f i c a n t l y l o w e r c a r b o h y d r a t e amount t h a n t h o s e f r o m m a t u r e wood, f o r t h e same t r a c h e i d l e n g t h . R e s u l t s a r e c o m p a r a b l e t o t h o s e r e p o r t e d by others:- b a s e d on g r o s s a n a l y s e s . 7. C o m p r e s s i o n wood t r a c h e i d s d i d n o t d i f f e r f r o m n o r m a l wood t r a c h e i d s i n . t h e amount o f h o l o c e l l u l o s e , f o r t h e same t r a c h e i d l e n g t h , b u t d i f f e r e d s i g n i f i c a n t l y i n , r e s p e c t t o a l p h a - c e l l u l o s e . I t was c o n c l u d e d t h a t c o m p r e s s i o n wood t r a c h e i d s may c o n t a i n l o w e r a l p h a - c e l l u l o s e t h a n m a t u r e (and normal) wood, b u t h i g h e r amount when compared t o j u v e n i l e wood, f o r t h e same t r a c h e i d l e n g t h . 8. S i g n i f i c a n t d i f f e r e n c e s i n m o r p h o l o g i c a l , phy- s i c a l , -and c h e m i c a l p r o p e r t i e s between e a r l y w o o d and l a t e w o o d o f c o n i f e r o u s s p e c i e s h a v e l o n g b e e n r e c o g n i z e d . E v i d e n c e o b - t a i n e d i n t h i s s t u d y shows t h a t , f o r t h e same t r a c h e i d l e n g t h , b o t h e a r l y w o o d and l a t e w o o d h a d s i m i l a r amounts o f a l p h a - c e l l u - l o s e , b u t d i f f e r i n g amounts o f h o l o c e l l u l o s e , l a t e w o o d t r a c h e i d s b e i n g h e a v i e r . 90. 9. 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 o f wood i n f e r - t i l i z e d t r e e s h a v e b e e n a t t r i b u t e d f o r m e r l y t o d i f f e r e n c e s i n growth r a t e and p e r c e n t a g e o f l a t e w o o d . E v i d e n c e was o b t a i n e d i n t h e p r e s e n t s t u d y t h a t , i n a d d i t i o n t o t h e above v a r i a b l e s , w e i g h t ' p e r u n i t l e n g t h o f c a r b o h y d r a t e t r a c h e i d s k e l e t o n s may a l s o c o n t r i b u t e t o an a l t e r a t i o n i n s p e c i f i c g r a v i t y . Changes' i n t r a c h e i d w e i g h t , a s s o c i a t e d w i t h f e r t i l i z a t i o n , f o r t h e most p a r t , were r e l a t e d t o t r a c h e i d l e n g t h . " Thus t h e b a s i c r e l a t i o n b etween l e n g t h and w e i g h t s t i l l e x i s t s i n f e r t i l i z e d wood; where t h e r e i s a r e v e r s i o n t o a p h a s e o f t e m p o r a r y j u v e n i l i t y , t r e e s p r o d u c e s h o r t e r t r a c h e i d s compared t o n o r m a l wood o f a compar- a b l e age.' The d a t a i n d i c a t e t h a t some t r e a t m e n t s may, however, g e n e r a t e l e s s a l p h a - c e l l u l o s e p e r u n i t l e n g t h when compared t o o t h e r s . T h i s d i f f e r e n c e i n b e h a v i o u r was s u g g e s t e d as a p o s s i b l e c a u s e f o r changes o b s e r v e d i n g r o s s wood s p e c i f i c g r a v i t y o f f e r t i l i z e d t r e e s . LITERATURE CITED B a i l e y , A . J . 1936. L i g n i n i n D o u g l a s f i r . C o m p o s i t i o n o f t h e m i d d l e l a m e l l a . I n d . E n g . Chem. ( A n a l . ) 8: 5 2 - 5 5 . B a i l e y , I.W. 19 20. The cambium and i t s d e r i v a t i v e t i s s u e s . I I . S i z e v a r i a t i o n s o f c a m b i a l i n i t i a l s i n gymno- sperm s and a n g i o s p e r m s . Am. J . B o t . 7: 3 5 5 - 3 6 7 . 1 9 2 3 . The cambium and i t s d e r i v a t i v e t i s s u e s . I V . The i n c r e a s e i n g i r t h o f t h e cambium. Am. J . B o t . 10: 4 9 9 - 5 0 9 . , and H.B. S h e p h a r d . 1915. S a n i o ' s l a w s f o r t h e v a r i a t i o n o f s i z e i n c o n i f e r o u s t r a c h e i d s . B o t a n . Gaz. 60: 6 6 - 7 1 . and A.F. F a u l l . 1934. The cambium and i t s d e r i v i t i v e t i s s u e s . . I X . S t r u c t u r a l v a r i a b i l i t y i n r e d w o o d ( S e q u o i a s e m p e r v i r e n s ) and i t s s i g n i f i c a n c e i n t h e i d e n t i f i c a t i o n o f f o s s i l woods. J . A r n o l d A r b o r e t u m 15: 233-254. B a n n a n , M.W. 1 9 6 5 . The l e n g t h , t a n g e n t i a l w i d t h , a nd l e n g t h / w i d t h r a t i o o f c o n i f e r t r a c h e i d s . C an. J . B o t . 43: 9 6 7-984. 1966. C e l l l e n g t h a n d r a t e o f a n t i c l i n a l d i v i s i o n s i n t h e cambium o f t h e s e q u o i a s . Can. J , B o t . 4 4 : 209-218. . 1967. A n t i c l i n a l d i v i s i o n s a n d c e l l l e n g t h i n c o n i f e r cambium. F o r . P r o d . J . 1 7 ( 6 ) : 63-69. 1970. A s u r v e y o f c e l l l e n g t h and f r e q u e n c y o f m u l t i p l i c a t i v e d i v i s i o n s i n t h e cambium o f c o n i f e r s C a n . J . B o t . 48: 1 5 8 5 - 1 5 8 9 . 10. a n d L L . B a y l y . 1956. 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D a d s w e l l . 1 9 52. The n a t u r e o f r e a c t i o n wood. I I I . C e l l d i v i s i o n a n d c e l l - w a l l f o r m a t i o n i n c o n i f e r s t e m s . A u s t r a l i a n J . S c i . R e s . S e r i e s B. B i o . S c i . 5: 3 8 5 - 3 9 8 . 1 0 3 . 147. W a r d r o p , A.B. and H.E. D a d s w e l l . 1 9 5 3 . The d e v e l o p m e n t o f t h e c o n i f e r t r a c h e i d . H o l z f o r s c h . 7: 33 - 3 9 . 148. . 1954. The d e v e l o p m e n t and s t r u c t u r e o f wood f i b r e s . P r o c . A P P I T A 8: 6-26. 149. W a t s o n , A . J . 1962. A s e m i m i c r o p r o c e d u r e f o r e s t i m a t i n g t h e r e s i s t a n t c a r b o h y d r a t e m a t e r i a l i n wood. T a p p i 45: 7 2 2-724. 150. and I.G. H o d d e r . 1954. R e l a t i o n s h i p b e t w e e n f i b r e s t r u c t u r e and h a n d s h e e t p r o p e r t i e s i n P i n u s t a e d a . P r o c . A P P I T A 8: 29 0 - 3 1 0 . 15 1 . W e s t i n g , A.H. • 1965. F o r m a t i o n a n d f u n c t i o n o f c o m p r e s - s i o n wood i n gymnosperms. B o t . Rev. 3 1 : 38 1 - 4 8 0 . 152. 1968. F o r m a t i o n and f u n c t i o n o f c o m p r e s - s i o n wood i n gymnosperms I I . B o t . Rev. 34: 51-78. 153. W e l l o n s , J.D. 1970. C e l l w a l l p o l y s a c c h a r i d e s i n s o u t h e r n p i n e w o o d - A r e v i e w . Wood S c i . 2: 247-254. 154. W e l l w o o d , R.W. 1 9 6 0 . S p e c i f i c g r a v i t y and t r a c h e i d l e n g t h v a r i a t i o n s i n s e c o n d - g r o w t h w e s t e r n h e m l o c k . J . F o r . 58: 3 6 1 - 3 6 8 . 155. . 1962. T e n s i l e t e s t i n g o f s m a l l wood s a m p l e s . P u l p P a p e r Mag. C a n . 63: T61-T67. 156. and J.W. W i l s o n . - 1965. The g r o w t h i n c r e - ment as a g u i d e t o p r o p e r t i e s i n c o n i f e r o u s woods. S e c t i o n 4 1 , IUFRO M e e t i n g , M e l b o u r n e , A u s t r a l i a . 157. , I f j u , G. and J.W. W i l s o n . 1 9 6 5 . I n t r a i n c r e m e n t p h y s i c a l p r o p e r t i e s i n c o n i f e r s . I n C e l l u l a r U l t r a s t r u c t u r e o f Woody P l a n t s ( e d . Cdt&). S y r a c u s e U n i v . P r e s s , S y r a c u s e , p p . 5 3 9 - 5 4 9 . 158. W h e e l e r , E. Y. ,• Z o b e l , B . J . and D.L. Weeks. 1966. T r a c h e i d l e n g t h and d i a m e t e r v a r i a t i o n i i u t h e b o l e o f l o b l o l l y p i n e . T a p p i 49: 48 4 - 4 9 0 . 159. W i l l i s , J.G. 1966. A D i c t i o n a r y o f F l o w e r i n g P l a n t s and F e r n s . C a m b r i d g e U n i v . P r e s s , C a m b r i d g e . 160. W i l s o n , J.W. 1970. P e r s o n a l c o m m u n i c a t i o n . 1 6 1 . and R.W. W e l l w o o d . 1 9 6 5 . I n t r a - i n c r e m e n t c h e m i c a l p r o p e r t i e s o f c e r t a i n w e s t e r n C a n a d i a n c o n i f e r o u s s p e c i e s . I n C e l l u l a r U l t r a s t r u c t u r e o f Woody P l a n t s ( e d . C6t£).. S y r a c u s e U n i v . P r e s s , S y r a c u s e , p p . 5 5 1 - 5 5 9 . 104. 162. W i s e , L . E . 1958. What i s wood c e l l u l o s e - a s e m a n t i c d i l e m m a . T a p p i 4 1 : 14A-22A. 163 . _____________ M u r p h y , M. a n d A.A. D ' A d d i e c o . 1946. C h l o r i t e h o l o c e l l u l o s e , i t s f r a c t i o n a t i o n a n d b e a r i n g o n s u m m a t i v e wood a n a l y s i s a n d on s t u d i e s on t h e h e m i - c e l l u l o s e s . P a p e r T r a d e J . 1 2 2 ( 2 ) : 3 5 - 4 3 . 164. W o d z i c k i , T. 1960. I n v e s t i g a t i o n on t h e k i n d o f L a r i x p o l o n i c a R a c . Wood f o r m e d u n d e r v a r i o u s p h o t o p e r i o d i c c o n d i t i o n s . - ' I . P l a n t s g r o w i n g i n n a t u r a l c o n d i t i o n s . A c t a S o c . B o t . P o l o n . 29: 713-730 ( F o r e s t r y A b s t r a c t s No. 4998. V o l . 2 2 ) . 165. W o r s t e r , H. a n d B.K. S u g i y a m a . 1962. The c a r b o h y d r a t e c o n t e n t and c o m p o s i t i o n o f some w e s t e r n woods r e l a t e d t o g r o w t h f a c t o r s . P u l p P a p e r Mag. C a n . 63: T395-T401. 166. Y i a n n o s , P.N. 1964.• The a p p a r e n t c e l l - w a l l d e n s i t y o f wood a n d p u l p f i b e r s . T a p p i 47: 4 6 8 - 4 7 1 . 167. Z i e g l e r , H. 1964. S t o r a g e , m o b i l i z a t i o n and d i s t r i b u t i o n o f r e s e r v e m a t e r i a l i n t r e e s . In The F o r m a t i o n o f Wood i n F o r e s t T r e e s ( e d . Zimmermann). A c a d e m i c P r e s s , New Y o r k . p p . 3 0 3 - 3 2 0 . 168. Zimmermann, M.H. ( e d . ) . 1964. The F o r m a t i o n o f Wood i n F o r e s t T r e e s . A c a d e m i c P r e s s , New Y o r k . 169. Z o b e l , B . J . 1 9 6 1 . I n h e r i t a n c e o f wood p r o p e r t i e s i n c o n i f e r s . S i l v a e G e n e t . 10: 6 5 - 7 0 . 170. and R.R. R h o d e s . 1 9 5 7 . S p e c i f i c g r a v i t y i n d i c e s f o r u s e i n b r e e d i n g l o b l o l l y p i n e . F o r . S c i . 3: 2 8 1 - 2 8 5 . 1 7 1 . .and R.L. M c E l w e e . 1958. V a r i a t i o n o f c e l l u - l o s e i n l o b l o l l y p i n e . T a p p i 41: 1 6 7 - 1 7 0 . 1 7 2 . , Thorb.j.ornsen, E. a n d F. H e n s o n . 1960. G e o g r a p h i c , s i t e and i n d i v i d u a l t r e e v a r i a t i o n i n wood p r o p e r t i e s o f l o b l o l l y p i n e . S i l v a e G e n e t . 9: 1 4 9 - 1 5 8 . 173 . , G o g g a n s , J . F . , M a k i , T.E. and F. H e n s o n . 1 9 6 1 . Some e f f e c t s o f f e r t i l i z e r s on wood p r o p e r t i e s o f l o b l o l l y p i n e . T a p p i 44: 1 8 6 - 1 9 2 . , TABLE I SINGLE TRACHEID (HOLOCELLULOSE SKELETON) WEIGHTS 9 , AND LENGTHS FOR NINE CONIFEROUS WOODS NO. OF TRACHEID TRACHEID LENGTH TRACHEID WEIGHT mm, ic ± 1 SD g SPECIES OBSERVATIONS DESIGNATION f x I O - 6 x ± 1 SD A r a u c a r i a c u n n i n g h a m i i 46 EW 8.12 + 0.73 3.43 + 0.75 N.S. LW 8.05 + 0.79 3.54 + 0.82 S e q u o i a s e m p e r v i r e n s 40 EW 6.15 + 0.50 1.66 + 0.42 * LW 6.73 + 0.64 2. 23 + 0.47 P i n u s l a m b e r t i a n a 35 EW 6.48 + 0.71 2.53 + 0.41 N.S. LW 6.15 + 1. 29 2.46 + 0.86 P o d o c a r p u s d a c r y d i o i d e s 28 EW 4.31 + 0.64 1. 40 + 0.33 * LW 4.97 + 0.57 1.72 + 0.29 P i c e a s i t c h e n s i s 25 EW 4.57 + 0.91 1.08 + 0.42 N.S. LW 4.54 + 0.90 1.08 + 0.40 Taxus b r e v i f o l i a 27 EW 2.10 + 0.32 0. 26 + 0.08 * LW 2.51 + 0.30 0.32 + 0.06 J u n i p e r u s v i r g i n i a n a 33 EW 2.20 + 0.47 0.19 + 0.08 * LW 2.62 + 0.36 0.26 + 0.06 C e p h a l o t a x u s w i l s o n i a n a 21 EW 1.95 + 0.17 0. 21 + 0.04 * LW 2.18 + 0.20 0. 28 + 0.05 P s e u d o t s u g a m e n z i e s i i 204 EW 4.35 + 1.65 1.07 + 0.67 ** LW 4.89 + 1.52 1.70 + 0.90 2 P s e u d o t s u g a m e n z i e s i i 48 EW 2.65 + 0.38 0.41 + 0.10 ** LW 3.03 + 0.36 0.66 + 0.18 P s e u d o t s u g a m e n z i e s i i 116 EW 2.92 + 0.56 0 . 50 + 0.22 ** 4 . . . P s e u d o t s u g a m e n z i e s i i LW 3.26 + 0.56 0.79 + 0.30 53 EW 2.64 + 0.22 0.42 + 0.14 ** LW 2.71 + 0.27 0.53 + 0.17 P s e u d o t s u g a m e n z i e s i i 52 EW 2.24 + 0.32 0.29 + 0.08 ** LW 2.45 + 0.27 0.40 + 0.11 P s e u d o t s u g a m e n z i e s i i 69 EW 2.57 + 0.24 0.32 + 0.11 ** LW 2.71 + 0.37 0. 47 + 0.13 Combined A v e r a g e 797 EW 3.72 + 1.82 0.90 + 0.89 ** LW 4.18 + 1.88 1.27 + 1.06 ( C o n t i n u e d ) 106. TABLE I ( C o n t i n u e d ) a A l l w e i g h t s e x p r e s s e d - i n o v e n d r y c o n d i t i o n EW E a r l y w o o d LW L a tewood x A v e r a g e l e n g t h o r w e i g h t SD S t a n d a r d d e v i a t i o n N.S. E a r l y w o o d and l a t e w o o d n o t s i g n i f i c a n t l y d i f f e r e n t a t t h e 5 % l e v e l ('t' t e s t ) * E a r l y w i r o d and l a t e w o o d s i g n i f i c a n t l y d i f f e r e n t a t t h e 5%. l e v e l ( 1 1 ' t e s t ) ** E a r l y w o o d and l a t e w o o d s i g n i f i c a n t l y d i f f e r e n t a t t h e 1% l e v e l ('t' t e s t ) G r oups w i t h i n D o u g l a s - f i r 1. B u t t l o g o f a 5 0 0 - y e a r - o l d t r e e ( I n c r e m e n t s 10, 20, 80, 150 300, 400)and I n c r e m e n t 20 f r o m g r o u p 3 b e l o w . 2. C o m p r e s s i o n wood - B r e a s t h e i g h t d i s k o f a 1 2 0 - y e a r - o l d t r e e , 3. NH4NO3 f e r t i l i z e d - B r e a s t h e i g h t d i s k o f a 2 9 - y e a r - o l d t r e e ( I n c r e m e n t s 21, 22, 23, 24, 25) f e r t i l i z e d a t ages 21 and 22. 4. U r e a f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t ; 2 4 - y e a r - o l d t r e e ( I n c r e m e n t s 15, 16) f e r t i l i z e d a t ages 13 and 15. 5. NPK f e r t i l i z e d -.12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t ; 2 4 - y e a r - o l d t r e e ( I n c r e m e n t s 15, 16) f e r t i l i z e d a t ages 13 and 15. 6. (NH4)2SO4 f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t , 2 4 - y e a r - o l d t r e e ( I n c r e m e n t s 15, 16) f e r t i l i z e d a t ages 13 and 15. TABLE I I SINGLE TRACHEID (ALPHA-CELLULOSE SKELETON) WEIGHTS 3 AND LENGTHS FOR NINE CONIFEROUS WOODS NO. OF TRACHEID TRACHEID LENGTH TRACHEID_WEIGHT OBSERVATIONS DESIGNATION mm, x ± 1 SD q x 10~6 x ± 1 SD A r a u c a r i a c u n n i n g h a m i i 33 EW 7.38 + 0.78 3.20 + 0. 53 N.S. LW 7.35 + 0.80 3.07 + 0.73 S e q u o i a s e m p e r v i r e n s 32 EW 5.36 + 0.81 1.58 + 0.48 ** LW 5.81 + 0.60 1.81 + 0. 42 P i n u s l a m b e r t i a n a 35 EW 5.37 + 0 . 62 1.92 + 0.37 ** LW 5.81 + 0.39 2.30 + 0.19 P o d o c a r p u s d a c r y d i o i d e s 43 EW 4.17 + 0.48 1.19 + 0. 24 N.S . LW 4.07 + 0.45 1.14 + 0. 29 P i c e a s i t c h e n s i s 46 EW 3.98 + 0.51 0.89 + 0. 24 N.S . LW 3.95 0.46 0.88 + 0.19 T a x u s b r e v i f o l i a 27 EW 1.93 + 0.39 0.18 + 0.06 N.S. LW 2.13 + 0.29 0. 23 + 0 .06 J u n i p e r u s v i r g i n i a n a 26 EW 1.87 + 0.24 0.19 + 0.05 * LW 2.12 + 0.27 0. 23 + 0.04 C e p h a l o t a x u s w i l s o n i a n a 21 EW 1.41 + 0.32 0.14 + 0.03 N.S . LW 1.64 + 0.27 0.16 + 0.04 P s e u d o t s u g a m e n z i e s i i 199 EW 3.33 + 1.21 0.95 + 0. 63 ** LW 3.88 + 1.27 1.26 + 0.74 P s e u d o t s u g a m e n z i e s i i 43 EW 2.31 + 0. 54 0.40 + 0.17 * LW 2.62 + 0.46 0.52 + 0.17 P s e u d o t s u g a m e n z i e s i i ; 112 EW 2.16 + 0.37 0.33 + 0.13 ** LW 2.70 + 0.54 0.62 + 0. 24 4 P s e u d o t s u g a m e n z i e s i i 42 EW 2.08 + 0.17 0.31 + 0.06 ** LW 2.39 + 0.23 0.43 + 0.09 P s e u d o t s u g a m e n z i e s i i 42 EW 1.99 + 0.30 0.29 . + 0 .07 * LW 2.18 + 0 .25 0.35 + 0.08 P s e u d o t s u g a m e n z i e s i i 46 EW 1.97 + 0.31 0.24 + 0.09 ** • LW 2.33 + 0.46 0.37 + 0.12 C o m b i n e d A v e r a g e 745 EW 3.12 + 1.55 0.79 + 0.77 ** LW 3.56 + 1.57 1.01 + 0. 84 ( C o n t i n u e d ) 108. TABLE I I ( C o n t i n u e d ) a A l l w e i g h t s e x p r e s s e d i n o v e n d r y c o n d i t i o n EW E a r l y w o o d LW L a t e w o o d x A v e r a g e l e n g t h o r w e i g h t SD S t a n d a r d d e v i a t i o n N.S. E a r l y w o o d a nd l a t e w o o d n o t s i g n i f i c a n t l y d i f f e r e n t a t t h e 5% l e v e l ( " f t e s t ) * E a r l y w o o d and l a t e w o o d s i g n i f i c a n t l y d i f f e r e n t a t t h e 5 % l e v e l ( ' t ' t e s t ) ** E a r l y w o o d a nd l a t e w o o d s i g n i f i c a n t l y d i f f e r e n t a t t h e 1% l e v e l ( ' t ' t e s t ) G r o u p s w i t h i n D o u g l a s - f i r 1. B u t t l o g o f a 5 0 0 - y e a r - o l d t r e e ( I n c r e m e n t s 10, 20, 80, 150 300, 4 0 0 ) a n d I n c r e m e n t 20 f r o m g r o u p 3 b e l o w . 2. C o m p r e s s i o n wood - B r e a s t h e i g h t d i s k o f a 1 2 0 - y e a r - o l d t r e e , 3. NH4NO3 f e r t i l i z e d - B r e a s t h e i g h t d i s k o f a 2 9 - y e a r - o l d t r e e ( I n c r e m e n t s 2 1 , 22, 23, 24, 25) f e r t i l i z e d a t ages 21 and 22. 4. U r e a f e r t i l i z e d - 12 mm i n c r e m e n t s c o r e f r o m b r e a s t h e i g h t ; 2 4 - y e a r - o l d t r e e ( I n c r e m e n t s 15, 16) f e r t i l i z e d a t ages 13 and 15. 5. NPK f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t ; 2 4 - y e a r - o l d t r e e ( I n c r e m e n t s 15, 16) f e r t i l i z e d a t ages 13 and 15. 6. (NH 4)2SC>4 f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t , 2 4 - y e a r - o l d t r e e ( I n c r e m e n t s 1 5 , 16) f e r t i l i z e d a t a g e s 13 and 15. TABLE I I I SINGLE TRACHEID (HOLOCELLULOSE SKELETON) WEIGHTS a AND LENGTHS FOR TWO DOUGLAS-FIR TREES INCREMENT NO. NO. OF TRACHEID TRACHEID LENGTH TRACHEID_WEIGHT TREE NO. FROM PITH OBSERVATIONS DESIGNATION mm, x ± 1 SD g x 1 0 ~ 6 x ± 1 SD 10 27 EW 2.17 + 0.30 0.24 + 0.07 LW 2.41 + 0.38 0.39 + 0.13 20 26 EW 3.58 + 0.70 0.85 + 0.35 LW 3.98 + 0.66 1.26 + 0.44 80 23 EW 5.61 + 0.70 1.65 + 0.52 LW 5.85 + 0.52 2.25 + 0.51 150 39 EW 5.97 + 0.60 1.77 + 0.47 LW 6.26 + 0.48 2.62 + 0.68 300 26 EW 5.79 + 0.69 1.63 + 0.45 LW 5.46 + 0.95 1.90 + 0.56 400 27 EW 5.85 + 0.61 1.42 + 0.28 LW 6.00 + 0.71 1.87 + 0.49 20 36 EW 2.92 + 0.59 0.56 + 0.21 LW 3.20 + 0.48 0.84 + 0.33 2 1 b 23 EW 2.92 + 0.57 0. 53 + 0.16 2 2 b LW 3.40 + 0.33 0.70 + 0.32 24 EW 2.93 + 0.45 0. 54 + 0.18 LW 3.22 + 0.54 0.74 + 0. 21 23 28 EW ' 2.98 + 0.48 0.44 + 0.12 LW 3.34 + 0.72 0. 85 + 0.36 24 15 EW 3.34 + 0.23 0.71 + 0.28 LW 3.58 + 0.27 1.02 + 0.32 25° 26 EW 2.67 + 0.73 0.37 + 0. 21 LW 2.86 + 0.52 0.62 + 0. 20 a b c T r e e T r e e A l l w e i g h t s e x p r e s s e d i n o v e n d r y c o n d i t i o n Y e a r s f e r t i l i z e d , C o m p r e s s i o n wood x A v e r a g e l e n g t h o r w e i g h t SD S t a n d a r d d e v i a t i o n EW E a r l y w o o d LW Latewood• 1 - B u t t l o g o f a 5 0 0 - y e a r - o l d t r e e ; 2 - NH 4N0 3 f e r t i l i z e d - B r e a s t h e i g h t d i s k o f a 2 9 - y e a r - o l d t r e e . TABLE IV SINGLE TRACHEID (ALPHA-CELLULOSE SKELETON) WEIGHTS 3 AND LENGTHS FOR TWO DOUGLAS-FIR TREES TREE -NO. INCREMENT NO. FROM PITH NO. OF OBSERVATIONS TRACHEID DESIGNATION TRACHEID LENGTH mm, x ± .1 SD TRACHEID_WEIGHT g x I O - 6 x ± 1 SD 10 28 EW 1.85 + 0.37 0.22 0.09 LW 2.18 + 0.33 0.31 + 0.12 20 34 EW 2.94 + 0.54 0.73 + 0.27 LW 3.46 + 0.62 0.96 + 0.32 80 24 EW 4.22 + 1.04 1.3 8 + 0.56 LW 4. 87 + 0.74 1.86 + 0.47 150 27 EW 4.26 + 1.22 1.50 + 0.70 LW 5.07 + 1.01 1.97 + 0.60 300 25 EW 4.-01 + 0.73 1. 24 + 0.37 LW 4.76 + 0.86 1.79 + 0.51 400 26 EW 4.48 + 0.70 1.33 + 0.44 LW 4 .54 + 0.94 1. 63 + 0.58 20 35 EW 2.39 + 0. 42 0.45 + 0.13 LW 2.80 + 0.42 0. 66 + 0. 21 2 1 b 22 EW 2.38 + 0.44 0.36 + 0.15 LW 2.79 + 0.47 0.64 + 0.22 22 b 20 EW 2.35 + 0.26 0.32 + 0.08 LW 2.66 + 0.28 0. 60 + 0.17 23 26 EW 2.03 + 0.19 0.31 + 0.07 LW 2.60 + 0.71 0.56 + 0.28 24 22 EW 2.12 + 0.33 0.38 . + 0.15 LW 2.98 + 0. 57 0.80 + 0. 27 2 5 C 22 EW 1.90 + 0.40 0. 27 + 0.16 LW •2.54 + 0.50 0.51 + 0.18 a A l l w e i g h t s e x p r e s s e d i n o v e n d r y c o n d i t i o n ? : b Y e a r s f e r t i l i z e d c C o m p r e s s i o n wood T r e e 1 - B u t t l o g o f a 5 0 0 - y e a r - o l d t r e e . T r e e 2 - NH 4N0 3 f e r t i l i z e d - B r e a s t h e i g h t d i s k o f a 2 9 - y e a r - o l d t r e e . x SD EW LW A v e r a g e l e n g t h o r w e i g h t S t a n d a r d d e v i a t i o n E a r l y w o o d L a t ewood TABLE V ANALYSIS MODEL (A) AND (B) RESULTS OF MULTIPLE CURVILINEAR COVARIANCE ANALYSES; DIFFERENCES AMONG SPECIES (HOLOCELLULOSE SKELETONS) (ADJUSTED FOR DIFFERENCES IN TRACHEID LENGTH) A) A n a l y s i s M o d e l . Group DF F T o t a l D i f f e r e n c e s f o r t e s t i n g s l o p e s Sums D i f f e r e n c e s f o r t e s t i n g l e v e l s Combined R e g r e s s i o n X B) 1 v s 2 vs 3 1 vs 2 vs 3 6 vs 7 v s 8 1 vs 2 vs 4 3 vs 5 vs 9 1, 2, 4, vs 6, 7, 8 vs ! 1 vs 2 3 4 5 7 8 v s 4 v s vs 5 Group 5 vs 9 1 1 1 6 6 6, 8 vs 4 v s 9 3 vs 1 v s vs v s v s vs v s S i g n i f i c a n c e i n S l o p e s L e v e l s 5 9 ** ** ** ** N.S , * N.S , ** ** * * ** * N.S, ** N.S , N.S , ** ** ** ** * ** ** ** * ** N.S. * ** N.S . ** ** N.S . ** Groups 1 A r a u c a r i a c u n n i n g h a m i i 2 S e q u o i a s e m p e r v i r e n s 3 P i n u s l a m b e r t i a n a . 4 P o d o c a r p u s d a c r y d i o i d e s 5 P i c e a s i t c h e n s i s 6 Taxus b r e v i f o l i a 7 J u n i p e r u s v i r g i n i a n a 8 C e p h a l o t a x u s w i l s o n i a n a 9 P s e u d o t s u g a m e n z i e s i i X - N.S. * ** I n d i c a t e s ' S i g n i f i c a n c e ' o r ' N o n - s i g n i f i c a n c e ' i n s l o p e s , I n d i c a t e s ' S i g n i f i c a n c e ' o r - ' N o n - s i g n i f i c a n c e 1 i n l e v e l s N o t s i g n i f i c a n t a t t h e 5% l e v S i g n i f i c a n t a t t h e 5% l e v e l S i g n i f i c a n t a t t h e 1% l e v e l 112. TABLE VI ANALYSIS MODEL (A) AND (B) RESULTS OF MULTIPLE CURVILINEAR COVARIANCE ANALYSES; DIFFERENCES AMONG SPECIES (ALPHA-CELLULOSE SKELETONS) (ADJUSTED FOR DIFFERENCES IN TRACHEID LENGTH) A) A n a l y s i s M o d e l Group DF F T o t a l D i f f e r e n c e s f o r t e s t i n g s l o p e s - X Sums D i f f e r e n c e s f o r t e s t i n g l e v e l s - Y Combined R e g r e s s i o n - B) Group S i g n i f i c a n c e i n S l o p e s L e v e l s 1 vs 2 v s 3 v s 4 vs 5 vs 9 ** ** 1 vs 2 v s 3 v s 5 N.S . ** 6 v s 7 v s 8 v s 9 ** ** 1 vs 2 vs 4 N.S. * 3 vs 5 vs 9 * ** 1, 2, 4 vs 9 ** ** 6, 7, 8 v s 9 ** * * 1 v s 2 N.S . N.S . 1 v s 3 N.S . ** 1 vs 4 N.S . N.S. 1 v s 5 N.S. ** 6 vs 7 * * 6 v s 8 N.S . ** 6, 8 vs 7 N.S . N.S. • 4 v s 9 ** ** 3 vs 5 N.S . ** 1 v s 9 ** ** Groups 1 A r a u c a r i a c u n n i n g h a m i i 2 S e q u o i a s e m p e r v i r e n s 3 P i n u s l a m b e r t i a n a 4 P o d o c a r p u s d a c r y d i o i d e s 5 P i c e a s i t c h e n s i s 6 Taxus b r e v i f o l i a 7 J u n i p e r u s v i r g i n i a n a 8 C e p h a l o t a x u s w i l s o n i a n a 9 P s e u d o t s u g a m e n z i e s i i X - I n d i c a t e s ' S i g n i f i c a n c e ' o r ' N o n - s i g n i f i c a n c e ' i n s l o p e s Y - I n d i c a t e s ' S i g n i f i c a n c e ' o r ' N o n - s i g n i f i c a n c e ' i n l e v e l s N.S. Not s i g n i f i c a n t a t t h e 5% l e v e l * S i g n i f i c a n t a t t h e 5% l e v e l ** S i g n i f i c a n t a t t h e 1% l e v e l 113. TABLE V I I MULTIPLE CURVILINEAR COVARIANCE ANALYSIS FOR TRACHEID WEIGHTS (HOLOCELLULOSE SKELETONS) IN JUVENILE, MATURE AND OVERMATURE WOOD OF A 500-YEAR-OLD DOUGLAS-FIR TREE (ADJUSTED FOR DIFFERENCES IN TRACHEID LENGTH) A Group DF F J u v e n i l e wood ( I n c r e m e n t " 1 0 ) 24 M a t u r e wood ( I n c r e m e n t s 20, 80, 150) 85 T o t a l : 109 D i f f e r e n c e s f o r t e s t i n g s l o p e s 2 1 .190 N. S. Sums: 111 D i f f e r e n c e f o r t e s t i n g l e v e l s 1 2 .010 N. S. Combined R e g r e s s i o n 112 B Group DF F J u v e n i l e wood ( I n c r e m e n t 10) 24 O v e r m a t u r e wood ( I n c r e m e n t s 300, 400) 50 T o t a l : 74 D i f f e r e n c e s f o r t e s t i n g s l o p e s 2 0 .722 N. s . Sums: 76 D i f f e r e n c e s f o r t e s t i n g l e v e l s 1 0 .355 N. s . Combined R e g r e s s i o n 77 C Group DF F M a t u r e wood ( I n c r e m e n t s 20, 80, 150) 85 O v e r m a t u r e wood ( I n c r e m e n t s 300, 400) 50 T o t a l : 135 D i f f e r e n c e s f o r t e s t i n g s l o p e s 2 7 .401** Sums: 137 D i f f e r e n c e s f o r t e s t i n g l e v e l s 1 29 .752** Combined R e g r e s s i o n 138 ——— —————————— — ——— — —————————— — ———— — — N.S. N o t s i g n i f i c a n t a t t h e 5% l e v e l ** S i g n i f i c a n t a t t h e 1% l e v e l 114. TABLE V I I I MULTIPLE CURVILINEAR COVARIANCE ANALYSIS FOR TRACHEID WEIGHTS (ALPHA-CELLULOSE SKELETONS) IN JUVENILE, MATURE AND OVERMATURE WOOD OF A 500-YEAR-OLD DOUGLAS-FIR TREE (ADJUSTED FOR DIFFERENCES IN TRACHEID LENGTH) A Group J u v e n i l e wood ( I n c r e m e n t 10) M a t u r e wood ( I n c r e m e n t s 20, 80, 150) T o t a l : DF 25 82 .107 F D i f f e r e n c e f o r t e s t i n g s l o p e s Sums: 2 109 2 .575 N. S. D i f f e r e n c e f o r t e s t i n g l e v e l s 1 7 .787 ** Combined R e g r e s s i o n 110 B Group J u v e n i l e wood ( I n c r e m e n t 10) O v e r m a t u r e wood ( I n c r e m e n t s 3 00, 400) T o t a l : DF 25 48 73 F D i f f e r e n c e f o r t e s t i n g s l o p e s Sums : 2 75 - 1 .413 N. S . D i f f e r e n c e f o r t e s t i n g l e v e l s 1 15 .069 ** Combined R e g r e s s i o n 76 C Group M a t u r e wood ( I n c r e m e n t s 20, 80, O v e r m a t u r e wood ( I n c r e m e n t s 3 00, 150) 400) T o t a l : DF 82 48 130 F D i f f e r e n c e f o r t e s t i n g s l o p e s Sums : 2 132 3 .898 * D i f f e r e n c e f o r t e s t i n g l e v e l s 1 0 .034 N. S . Combined R e g r e s s i o n 133 N.S. N o t s i g n i f i c a n t a t t h e 5% l e v e l * S i g n i f i c a n t a t t h e 5% l e v e l ** S i g n i f i c a n t a t t h e 1% l e v e l 115. TABLE IX ANALYSIS OF COVARIANCE AND ADJUSTED MEAN VALUES OF EARLYWOOD AND LATEWOOD TRACHEID WEIGHTS a IN SIX DOUGLAS-FIR TREES (ADJUSTED FOR DIFFERENCES IN TRACHEID LENGTH). HOLOCELLULOSE ALPHA-CELLULOSE SKELETONS SKELETONS S o u r c e DF F DF F T r e e s 5 1.23 N.S. 5 2.45* E a r l y w o o d / L a t e w o o d 1 97.92 ** 1 0.00 N.S, E r r o r 534 475 A d j u s t e d Mean T r a c h e i d S k e l e t o n W e i g h t s E a r l y w o o d 0.7 5 0.70 Latewood 0.94 0.7 0 — 6 a A l l w e i g h t s e x p r e s s e d as g x 10 i n o v e n d r y c o n d i t i o n N.S. N o t s i g n i f i c a n t a t t h e 5% l e v e l * S i g n i f i c a n t a t t h e 5% l e v e l ** S i g n i f i c a n t a t t h e 1% l e v e l 116. TABLE X ANALYSIS OF COVARIANCE AND ADJUSTED MEAN VALUES OF EARLYWOOD AND LATEWOOD TRACHEID WEIGHTS a IN NINEb CONIFEROUS SPECIES (ADJUSTED FOR DIFFERENCES IN TRACHEID LENGTH) HOLOCELLULOSE SKELETONS ALPHA-CELLULOSE SKELETONS S o u r c e DF Groups 13 20.91 ** E a r l y w o o d / L a t e w o o d 1 65.43 ** E r r o r 781 DF 13 1 729 64.66 ** 1.88 N.S A d j u s t e d Mean T r a c h e i d S k e l e t o n W e i g h t s E a r l y w o o d L atewood 1.02 1.17 0.90 0.91 a A l l w e i g h t s e x p r e s s e d as g x 10 i n o v e n d r y c o n d i t i o n b , c I n c l u d e s a l l s p e c i e s and t h e s i x t r e e s w i t h i n D o u g l a s - f i r ; N.S. N o t s i g n i f i c a n t a t t h e 5% l e v e l ** S i g n i f i c a n t a t t h e 1% l e v e l TABLE XI SELECTED WOOD AND TRACHEID PROPERTIES OF THREE F E R T I L I Z E D DOUGLAS-FIR TREES C h a r a c t e r i s t i c s T r e e 84 T r e e 90 T r e e 74 T r e a t m e n t C o m p o s i t i o n U r e a NPK ( N H 4 ) 2 S O 4 I n c r e m e n t f r o m p i t h 12 15* 16 12 15* 16 12 15* 16 S p e c i f i c g r a v i t y l 0.414 0.430 0.436 0.403 0.374 0.352 0.381 0.371 0. 337 ( 3 - y e a r age segment) P e r c e n t a g e o f l a t e w o o d 28.6 23.1 20.0 24.7 18.6 18.6 24. 6 14.3 10. 7 I n c r e m e n t w i d t h , mm 4.2 6.5 5.5 5.1 10.5 10.7 5.7 7.7 7. 5 T r a c h e i d l e n g t h , mm 2.26 3.30 2.80 2.10 2.36 2.55 2.20 2. 26 2. 40 T a n g e n t i a l t r a c h e i d 27.0 28.0 27.5 29 . 8 29.5 29 .4 29.2 27. 8 26. 2 d i a m e t e r , m i c r o n R a d i a l t r a c h e i d 34.5 43.1 40.0 33.0 38.0 40.5 38.0 40.5 46. 0 d i a m e t e r , m i c r o n T a n g e n t i a l d o u b l e 7.6 7.5 7.0 7.5 7.5 7.5 7.0 7.5 8. 0 ,-;.wall t h i c k n e s s , m i c r o n R a d i a l d o u b l e w a l l 9.0 10.0 10.5 11.0 12.0 12.5 11.5 10. 2 10. 5 t h i c k n e s s , m i c r o n * F i r s t f e r t i l i z e d a t t h e age o f 13 and r e f e r t i l i z e d a t age 15 a t t h e r a t e of 200 lb Cpounds per acre of N, P 2 ^ § ' o r K 2 ^ e < 3 u i v a ^ - e n t s ) 1. Maximum m o i s t u r e c o n t e n t method. 118. TABLE X I I RESULTS OF MULTIPLE CURVILINEAR COVARIANCE ANALYSES; DIFFERENCES WITHIN SPECIES IN TREATED AND UNTREATED DOUGLAS-FIR (ADJUSTED FOR DIFFERENCES IN TRACHEID LENGTH) A. (HOLOCELLULOSE (ALPHA-CELLULOSE SKELETONS) SKELETONS) G r o u p a DF F DF F 1 201 196 3 113 109 4 50 39 5 49 39 6 66 42 T o t a l 479 425 D i f f e r e n c e s f o r t e s t i n g s l o p e s 8 1.915 N .S. 8 5 .650 ** Sums 487 433 D i f f e r e n c e s f o r t e s t i n g l e v e l s 4 1.115 N .S . 4 7 .405 ** Combined R e g r e s s i o n 491 437 B. S i g n i f i c a n c e i n S i g n i f i c a n c e i n Group S l o p e s L e v e l s S l o p e s L e v e l s 4 v s 5 v s 6 N.S. ** * * ** 4 v s 6 N.S. * * ** ** 4 v s 5 N.S . N.S . * * N.S . 3 vs 4 N.S . * N.S . N.S . 4 vs 7 N.S . N.S . N.S . ** 5 vs 7 N.S . N.S . N.S. ** 6 v s 7 N.S . N.S . N.S. N.S . 3 vs 7 ** N.S . ** N.S . N.S. N o t s i g n i f i c a n t . a t t h e 5% l e v e l * S i g n i f i c a n t a t t h e 5% l e v e l ** S i g n i f i c a n t a t t h e 1% l e v e l a Group 2 - C o m p r e s s i o n wood o f D o u g l a s - f i r i s u n d e r s e p a r a t e h e a d i n g ( C o n t i n u e d ) 119. TABLE X I I ( C o n t i n u e d ) Groups w i t h i n D o u g l a s - f i r 1. B u t t l o g o f a 5 0 0 - y e a r - o l d t r e e ( I n c r e m e n t s 10, 20, 80, 150, 300, 400) and I n c r e m e n t 20 f r o m g r o u p 3 b e l o w . ? 3. NH4NO3 f e r t i l i z e d - B r e a s t h e i g h t d i s k o f a 2 9 - y e a r - o l d t r e e (Increments,.'21, 22, 23, 24, 25) f e r t i l i z e d a t ages 21 and 22. 4. U r e a f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t ; 2 4 - y e a r - o l d t r e e ( I n c r e m e n t s 15, 16) f e r t i l i z e d a t ages 13 and 15. 5. NPK f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t ; 2 4 - y e a r - o l d t r e e ( I n c r e m e n t s 15, 16) f e r t i l i z e d a t ages 13 and 15. 6. (NH4)2SC>4 f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t ; 2 4 - y e a r - o l d t r e e ( I n c r e m e n t s 15, 16) f e r t i l i z e d a t ages 13 and 15. 7. J u v e n i l e wood - I n c r e m e n t 10 f r o m Group 1 a b o v e . 120. FOREWORD TO FIGURES F i g u r e s 1 t o 10, 13, 15 t o 1 8 - - P l o t t i n g and c u r v e f i t t i n g by c o m p u t e r . M a t h e m a t i c a l m odel f o r c u r v e f i t t i n g : Y = b Q + b± X + b 2 X 2 _ g Where Y = Ovendry w e i g h t o f t r a c h e i d i n g x 10 X = T r a c h e i d l e n g t h i n mm - g r e e n c o n d i t i o n F i g u r e 1 1 — P l o t t i n g and c u r v e f i t t i n g by c o m p u t e r . The l i n e o f b e s t f i t c o n d i t i o n e d t o p a s s t h r o u g h t h e o r i g i n , I n a l l f i g u r e s , t h e m a t h e m a t i c a l m o d els g i v e n a r e t h e e q u a t i o n s o f b e s t f i t . F i g u r e 1: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s f r o m A r a u c a r i a c u n n i n g h a m i i . H I I I 1 1 1 1 1 1 1 3.0 0.98 1.96 2.94 3.92 4.90 5.98 6.86 7.B4 8.92 9 BC Tracheid Length, mm' F i g u r e 2: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s f r o m S e q u o i a s e m p e r v i r e n s . H0LO-SP2 122, A. -2.263 + 0.6580K r2 = 0,64, SE = 0.3OS; DF = 38 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. x IO"6 X = TRACHEID LENGTH IN ffl - GREN CONDITION l'.96 — I 2.94 - 1 3.92 . 4.9© 5.88 6.86 ALPHA SP2 B, -0.1590 + O.OStX2 •-2 = 0.93; SE = 0.19; DF = 30 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. x 10-6 X = TRACHEID LENGTH IN HI - GREN CONDITION —1 3.98 -r 2.94 3.92 4 . 9 0 J . f l f l Tracheid Length,mm- F i g u r e 3: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s f r o m P i n u s l a m b e r t i a n a . 123. A. 88 H0L0-SP3 Y = -1.1278 + 0.622X r.2 = 0.93; SE = 0.183; DF = 3 WHERE Y = OVENDRY WEIGHT CF TRACHEID IN G. x 10-6 X = TRACHEID LENGTH IN W - GREN CONDITION —I 0.90 —I 2.54 —1 3.92 —1 4.90 —1 E.B6 -1 T.B4 ~1 8.82 RLPHfi SP3 B. -1.080 + 0.572X r2 = 0.860; SE = 0.130; DF = 3 WHERE Y = OVENDRY WEIGHT CF TRACHEID IN G. x IO"6 X = TRACHEID LENGTH IN Wi - GREN CONDITION - r - — 2.94 3.92 4.9<? Tracheid Length, mm- ~T ?.8e F i g u r e 4: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s f r o m P o d o c a r p u s d a c r y d i o i d e s . H0L0-SP4 Y = 0 .019 + 0.0t97X2 r'2 = 0.929- SE = 0.109; DF = 26 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. X Iff* X = TRACHEID LENGTH IN HI - GREN CONDITION 124 - 1 — 0.96 I 2.94 I 3.92 -1 5.88 - 1 6.86 —1 7.64 —1 8.82 (£> 0-.0 RLPHfi SP4 B. Y = 0.0101 + 0.0527X2 r2 = 0.90; SE = 0.068; DF = 1 WHERE: Y = OVENDRY WEIGHT OF TRACHEID IN G. x 10-6 X = TRACHEID LENGTH IN MM - GREEN CONDITION I 1.96 T — I 2.94 9.92 4.90 Tracheid Length, mm- -r - 1 — 7.84 F i g u r e 5: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s f r o m P i c e a . s i t c h e n s i s . H0L0-SP5 Y = -0.7389 + 0.H939X r 2 = 0.948 S E E = 0.080; DF = 23 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. x 10-6 X = TRACHEID LENGTH IN IT! - GREEN CONDITION — i — 0.9fl -1 1.96 —1 2.94 -1 3.92 —1 r 4.90 5.6 -1 6.86 ? 6.82 ALPHA SP5 Y = -0.0056 + 0, 1-2 = 0.920; S E E = 0.076; DF = W WHERE Y = OVENDRY WEIGHT CF TRACHEID IH G, x 10-6 X = TRACHEID LENGTH IN m - GREEN CONDITION — I — 2.94 -i r 4.9C 5.B 3.32 Tracheid Length, mm F i g u r e 6: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s from Taxus b r e v i f o l i a . 40L0-SP6 Y = -0.15)1 + 0.1917X r 2 = 0.833; S__ = 0.033; DF = 25 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. x 10-6 X = TRACHEID LENGTH IN ffl - GREEN CONDITION RLPHq SP6 Y = 0.0155 + O.COW i 2 = 0.927; S_E = 0,017; DF = 25 WHEE Y = OVENDRY WEIGHT OF TRACHEID IN G, x 10-6 X = TRACHEID LENGTH IN Ml - GREEN CONDITION —I— 2.94 -1 1 3.92 4.9« Tracheid Length,mm- — I 6.96 F i g u r e 7: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s f r o m J u n i p e r u s v i r g i n i a n a . 127. HGL0-SP7 Y = 0,010 + 0.0347X2 r2 = 0.894; SE = 0.025; DF = 31 WHERE Y = OVENDRY WEIGHT OF TRACHEID ] G. x IO"6 X = TRACHEID LENGTH IN PM - GREN CONDITION -1 1.98 —I 3.92 - 1 5.BB —I G.B6 - 1 T.B4 ? a.82 9.Bp flLPHfi SP7 = -0.102 + 0.1549X r2 = 0.918; SE = 0.013; DF = 24 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. x 10-6 X = TRACHEID LENGTH IN W - GREK CONDITION —I 2.94 3.92 4.90 Tracheid Length, mm F i g u r e 8: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s f r o m C e p h a l o t a x u s w i l s o n i a n a . 128 . H0L0-SP8 A. Y = -0.2790 + 0.2526X r 2 = 0.921; SEE = 0.016; DF = 19 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. x 10-6 X = TRACHEID LENGTH IN m - GREEN CONDITION - 1 0.98 -1 1.96 -1 2.94 -1 3.92 RLPHfi SP8 B. Y = 0,0618 + 0.035GX2 r 2 = 0.818; SEE = 0.013; DF = 19 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G, x 10-6 X = TRACHEID LENGTH IN FN - GREEN CONDITION 8 ^ I I I I I 1 1 1 1 3.98 1.66 2.94 3.92 4.90 5.88 6.96 7.84 8.82 Q.80 Tracheid Length,mm- F i g u r e 9: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r samples f r o m P s e u d o t s u g a m e n z i e s i i . 129. 3 8 4 A. 3 2 0 o Co nd il >. 2-56 ,e nd i o i o 1-92 X cn a> 1-28 13 oc he i H 0 - 6 4 0-00"— 000 y = 00769+00554x 2 r 2 =0-843 3EE =0-342 DF=202 Where y = oven dry weight of tracheid in g xlO' x = tracheid length in mm -green condition least-squares fitted line 0-98 1-96 2-94 3-92 490 5-88 Tracheid Length, mm 6-86 7-84 8-82 B. 4-48 B 3-84 o o fT 3 20 T3 O 2 56 •92 1-28 - _ a> u £ 0-64 000 y = -0-3189+0-2184 x +00437 x 2 R 2 =0-986 S E E =0-083 DF = I96 Where y = oven dry weight of tracheid in g xlO""6 x= tracheid length in mm -green condition least-squares fitted line 000 0-98 2-94 3-92 4-90 5-88 6-86 7-84 Tracheid Length,mm- F i g u r e 10: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s f r o m n i n e c o n i f e r o u s s p e c i e s . ( L e a s t - s q u a r e s f i t t e d l i n e ) . HOLO SP1-9 130 Y = -0.1463 + 0.099OX + 0.0436X2 fi2 = 0.919; SEE = 0.319; DF =456 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G, X 10"6 X = TRACHEID LENGTH IN MM - GREEN CONDITION Tracheid Length, mm F i g u r e 11: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r s a m p l e s f r o m n i n e c o n i f e r o u s s p e c i e s . (The l i n e o f b e s t f i t c o n d i t i o n e d t o p a s s t h r o u g h t h e o r i g i n ) . 131 HOLD SP1-9 - 0.03E8X + O.OWX2 R2 = 0,919, SE = 0,320; DF = 457 WHERE Y = OVENDRY WEIGHT CF TRACHEID IN G. x 10-6 X = TRACHEID LENGTH IN ffl - GREN CONDITION 1 1 r 1 1 1 r a.a 0 98 1.96 i.94 3 9 i 4.9o 5.88 B.I — I — T.B4 ALPHA SP1-9 = 0.0758X + O.OWiX2 R2 = 0.952; SE> 0.196; DF=459 WERE Y = OVENDRY '-.'EIGHT OF TRACHEID IN G, x 10-E X = TRA.CHEID LENGTH IN m - GREN CONDITION -r —r — i 8.82 2.94 3.92 4.9C Tracheid Length,mm- F i g u r e 12: T r a c h e i d w e i g h t , mean t r a c h e i d l e n g t h and s p e c i f i c g r a v i t y p a t t e r n s a c r o s s t h e b u t t l o g o f a 5 0 0 - y e a r ^ o l d D o u g l a s - f i r t r e e . ( E a c h p o i n t r e p r e s e n t s t h e a v e r a g e t r a c h e i d w e i g h t o r l e n g t h f o r t h e i n c r e m e n t . S p e c i f i c g r a v i t y v a l u e s f r o m Kennedy a n d W a r r e n ( 7 0 ) ) . • _ £ T A F i g u r e 13: R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h f o r compres- s i o n wood and n o r m a l wood o f D o u g l a s - f i r . HOLO COMP WOOD 133 A. MULTIPLE CURVILINEAR COVARIANCE ANALYSES CROUP DE 201 15 216 2 218 1 219 DOUGLAS-FIR (NORMAL WOOD) COMPRESSION WOOD TOTAL DIFFERENCE FOR TESTING SLOPES SUMS DIFFERENCE FOR TESTING LEVELS COMBINED REGRESSION N.S, NOT SIGNIFICANT AT THE 5? LEVEL 0.287N.S. 0.C60N.S, Y = -0.5959 + 0.1012X r 2= 0.717; SE = 0.103 DF = 16 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. x 10-6 X = TRACHEID LENGTH IN MM - GREEN CONDITION ° s , least-squares fitted line ^ f o r Douglas-fir least-squares fitted line for Douglas-fir Compression Wood —I 2.94 RLPHP, COMP WOOD MULTIPLE CURVILINEAR COVARIANCE ANALYSES B, GROUP DOUGLAS-FIR (NORMAL WOOD) COMPRESSION WOOD TOTAL DIFFERENCE FOR TESTING SLOPES SUMS DIFFERENCE FOR TESTING LEVELS COMBINED REGRESSION ** SIGNIFICANT AT THE 11 LEVEL DE 196 10 236 2 8.760" 1 7.613*' 239 Y = 0.021 + 0.C v2 = 0.90; SE = 0.057 DF = 1 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G, X 10-6 X = TRACHEID LENGTH IN MM - GREEN CONDITION least-squares fitted line for Douglas-fir least-squares fitted line for Douglas-fir Compression Wood — I — 1.96 -| 5.88 3.92 4.90 Tracheid Length, mm F i g u r e 14: T r a c h e i d w e i g h t , i n c r e m e n t w i d t h and mean t r a c h e i d l e n g t h p a t t e r n s a c r o s s s i x c o n - s e c u t i v e i n c r e m e n t s o f a f e r t i l i z e d (NH4NO3) D o u g l a s - f i r w h i c h p r o d u c e d com- p r e s s i o n wood 4 . y e a r s a f t e r f i r s t f e r t i l i - z a t i o n . ( E a c h p o i n t r e p r e s e n t s t h e a v e r a g e t r a c h e i d w e i g h t o r l e n g t h f o r t h e i n c r e m e n t ) . Tracheid Weight, g- x l O - 6 (Ovendry Condition) Increment Width, mm- Mean Tracheid Length, mm- F i g u r e 15: " R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h i n U r e a t r e a t e d and n o r m a l D o u g l a s - f i r . 135. A. ion  •o s c o- 0  >. •o > 0 O 0 g •v X We ic to •5 <u f 0 r— B, HOLQ UREA MULTIPLE CURVILINEAR COVARIANCE ANALYSES GROUP DF F DOUGLAS-FIR 201 UREA FERTILIZED DOUGLAS-FIR 50 TOTAL 251 DIFFERENCES FOR TESTING SLOPES 2 0.948 N.S. SIMS 253 DIFFERENCES FOR TESTING LEVELS 1 0.08 N.S. COMBINED REGRESSION 254 N.S. NOT SIGNIFICANT AT THE 5% LEVEL Y = -0.2423 + 0.097X2 r2 = 0.635; SE - 0.103 DF = 51 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN e. x 10-6 X = TRACHEID LENGTH IN MM - GREEN CONDITION <r least-squares fitted line for Douglos-fir ^least-squares fitted line for Douglas-fir, Urea fertilized —I 1 1 1.96 2.94 3.92 —I 4.90 I 1 5.88 6.86 1.82 9.8c ALPHA UREA MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP DF F DOUGLAS-FIR 196 UREA FERTILIZED DOUGLAS-FIR 39 TOTAL 235 DIFFERENCES FOR TESTING SLOPES 2 0.721 N.S, SUMS 237 DIFFERENCES FOR TESTING LEVELS 1 2,517 N.S, COMBINED REGRESSION 238 N.S. NOT SIGNIFICANT AT THE 5Z LEVEL Y = 0.6502 - 0.6410X+ 0.264X2 R.2 = 0.942; SE = 0.025 DF = 39 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G, X 10"6 X - TRACHEID LEN3TH IN Ml - GREEN CONDITION least-squares fitted line for Douglas-fir ~leost-squares fitted line for Douglas-fir.Urea fertilized T 2.94 3.92 4.90 Tracheid Length,mm- , A F i g u r e 16: R e l a t i o n s h i p b e t w e e n t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h i n NPK t r e a t e d and n o r m a l D o u g l a s - f i r . HOLO NPK MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP DOUGLAS-FIR NPK FERTILIZED DOUGLAS-FIR TOTAL DIFFERENCES FOR TESTING SLOPES SUMS DIFFERENCES FOR TESTING LEVELS COMBINED REGRESSION DF F 201 19 250 2 0.106.N.S, 252 1 0.581 N.S. 253 Y = -0.017 + 0.069W2 r2 = 0.710.- SE = 0.063 DF = 50 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. X 10-6 X = TRACHEID LENGTH IN MM - GREEN CONDITION N,S. NOT SIGNIFICANT AT THE 5? LEVEL ALPHA NPK MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP DF DOUGLAS-FIR 196 NPK FERTILIZED DOUGLAS-FIR 39 TOTAL 235 DIFFERENCES FOR TESTING SLOPES 2 SUMS 237 DIFFERENCES FOR TESTING LEVELS 1 COMBINED REGRESSION 238 " SIGNIFICANT AT THE V. LEVEL N.S. NOT SIGNIFICANT AT THE 5% LEVEL 1.753 N.S. = 0.030 Y = Q035 + 0.C <2 = 0,861; SE DF = 10 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. X IO"6 X = TRACHEID LENGTH IN MM - GREEN CONDITION Tracheid Length, mm- »A F i g u r e 17: R e l a t i o n s h i p between t r a c h e i d weight ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) skeletons) and l e n g t h i n (NH4)2S04 t r e a t e d and normal D o u g l a s - f i r . HOLO NS04 Y = -0.0758 + 0.0676X2 '2 = 0,687 SE = 0.079 DF = 67 WHERE Y = OVENDRY WEIGHT OF. TRACHEID IN G, x 10-6 X = TRACHEID LENGTH IN MM - GREEN CONDITION ALPHA NS04 MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP DF F DOUGLAS-FIR 201 (NHi|)2S0i( FERTILIZED DOUGLAS FIR 6 TOTAL 267 DIFFERENCES FOR TESTING SLOPES 2 0.309 N.S SUMS 269 DIFFERENCES FOR TESTING LEVELS 1 1.61)5 N.S COMBINED REGRESSION 270 N.S. NOT SIGNIFICANT AT THE 57. LEVEL MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP DF F DOUGLAS-FIR 196 (NhtyjSQi) FERTILIZED DOUSLAS-FIR 12 TOTAL 238 DIFFERENCES FOR TESTING SLOPES 2 13,631 SUMS 210 DIFFERENCES FOR TESTING LEVELS 1 21,103* COMBINED REGRESSION 211 Y = -0,253 + 0.259X ' <2 = 0.815,- SEE - 0.019 DF = 13 WHERE Y = OVENDRY WEIGHT OF TRACHEII IN G. X 10-6 X " TRACHEID LENGTH IN MM - GREEN CONDITION ' SIGNIFICANT AT THE 1% LEVEL Tracheid Length,mm- . A F i g u r e 18: , R e l a t i o n s h i p between t r a c h e i d w e i g h t ( h o l o c e l l u l o s e (A) and a l p h a - c e l l u l o s e (B) s k e l e t o n s ) and l e n g t h i n NH4NO3 t r e a t e d and n o r m a l D o u g l a s - f i r . HOLD HRNET 2-6 MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP jf p DOUGLAS-FIR 201 NH4NO3 FERTILIZED DOUGLAS FIR 113 TOTAL 311 DIFFERENCES FOR TESTING SLOPES 2 56,192*' SUMS 316 DIFFERENCES FOR TESTING LEVELS 1 11,581" COMBINED REGRESSION 317 ** SIGNIFICANT AT THE 1% LEVEL Y = 0.6561 - 0.5187X + 0.1625 X2 R2 = 0.781; SE E = 0.113; DF = 149 WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. x IO - 6 X = TRACHEID LENGTH IN Ml - GREEN CONDITION least-squares fitted line for Dauglas-fir •least-squares fitted line for Douglas-fir, N H 4 N O 3 fertilized 1.96 2-94 - r 6.86 —1 8.82 flLPHR HRNEY 2-6 MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP DF F DOUGLAS FIR 196 NH1IO3 FERTILIZED DOUGLAS-FIR 109 TOTAL 305 DIFFERENCES FOR TESTING SLOPES 2 51,519" SUMS 307 DIFFERENCES FOR TESTING LEVELS 1 2.395 N.S. COMBINED REGRESSION 308 N.S. NOT SIGNIFICANT AT THE 57. LEVEL ** SIGNIFICANT AT THE 12 LEVEL Y = -0.0507 + 0.085D<2 T2 = 0.909; S E E = 0,072; DF = 115 WHERE Y = OVENDRY WEIGHT CF TRACHEID IN G, x 10-6 X = TRACHEID LENGTH IN Ml - GREEN CONDITION least-squares fitted line for Douglas-fir leost-squores filled line for Douglas-fir, N H 4 N O 3 fertilized 3.92 4.90 Tracheid Length, mm- —I 6.86 A P P E N D I C E S APPENDIX I DESCRIPTION OF GROWTH INCREMENTS INCLUDED IN THE STUDY S p e c i e s Age From P i t h Y r . Wood Zone I n c r e m e n t W i d t h mm L a t e w o o d W i d t h mm M e a s u r e d T r a c h e i d L e n g t h ct A v e r a g e nun Range - A r a u c a r i a c u n n i n g h a m i i — H 2.4 0.4 7.42 5.10-9.79 S e q u o i a s e r a p e r v i r e n s - H 2.5 0.5 5.97 • 4.20-8.50 P i n u s l a m b e r t i a n a - H 1.4 0.3 6.29 3.75-8.12 P o d o c a r p u s d a c r y d i o i d e s - H 3.6 0.6 4.59 2.92-6.04 P i c e a s i t c h e n s i s 36 H 4.4 1.4 4.70 2.92-5.84 Taxus b r e v i f o l i a 70 H 2.3 0.6 2.14 1.44-2.96 J u n i p e r u s v i r g i n i a n a - H 3.4 0.8 2.37 1.42-2.99 C e p h a l o t a x u s w i l s o n i a n a 8 S 6.5 0.8 2.13 1.70-2.56 ^ P s e u d o t s u g a m e n z i e s i i 10 H 2.0 0.9 2.37 1.88-2.71 20 H 5.8 2.4 3.97 2.14-4.55 80 H 3.4 1.4 5.46 3.96-6.67 150 H 1.7 0.7 5.92 3.33-7.08 300 H 1.0 0.3 4.96 3.13-6.88 2 400 H 0.7 0.2 5.89 3.45-7.08 P s e u d o t s u g a m e n z i e s i i 73 H 4.9 3.5 2.84 0.85-3.27 2 83 S 5.0 3.2 2.89 1.99-3.27 P s e u d o t s u g a m e n z i e s i i 20 H 5.9 2.5 2.92 1.20-3.80 21 H 7.7 3.7 2.50 1.45-4.20 22 H 10.0 3.3 2.51 1.65-3.95 23 H 8.5 2.4 2.40 1.90-4.20 24 S 8.5 2.6 2.81 1.65-3.77 25 S 8.6 3.7 2.30 1.05-3.85 P s e u d o t s u g a m e n z i e s i i 15 S 6.5 1.5 3.31 2.70-3.98 16 S 5.5 1.1 2.80 2.13-3.56 P s e u d o t s u g a m e n z i e s i i 15 H 10. 5 2.0 2.39 1.85-3.13 16 H 10.7 2.0 2.54 1.99-3.13 P s e u d o t s u g a m e n z i e s i i 15 H 7.7 1.1 2.23 1.42-2.99 16 H 7.5 0.9 2.39 1.99-3.56 ( C o n t i n u e d ) 140. APPENDIX I ( C o n t i n u e d ) A v e r a g e b a s e d on a minimum o f 50 t r a c h e i d s H e a r t w o o d Sapwood B u t t , l o g o f a 5 0 0 - y e a r - o l d t r e e C o m p r e s s i o n wood - B r e a s t h e i g h t d i s k o f a 1 2 0 - y e a r - o l d t r e e NH4NCU f e r t i l i z e d - B r e a s t h e i g h t d i s k o f a 2 9 - y e a r - o l d t r e e j f e r t i l i z e d a t ages 21 and 22 (500 l b / a c r e ) U r e a f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t , 2 4 - y e a r - o l d t r e e ; f e r t i l i z e d a t ages 13 and 15 (200 l b / a c r e o f N,P 20 p r K 2 0 e q u i - v a l e n t s ) NPK f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t , 2 4 - y e a r - o l d t r e e ; f e r t i l i z e d a t ages 13 and 15 (200 l b / a c r e o f N^-O^orK.O e q u i - v a l e n t s ) (NH^) 2SO^ f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t , 2 4 - y e a r - o l d t r e e ; f e r t i l i z e d a t ages 13 and 15 (200 l b / a c r e o f N , P 2 0 5 o r K 2 0 e q u i v a l e n t s ) APPENDIX I I SUMMARY OF HOLOCELLULOSE AND ALPHA-CELLULOSE. YIELDS AND ESTIMATES OF LIGNIN IN SOME OF THE SAMPLES STUDIED (BASED ON OVENDRY, EXTRACTIVE-FREE WOOD) D e s c r i p t i o n I n c r e m e n t P o r t i o n No. o f H o l o c e l l u - A l p h a - C e l l u - S p e c i e s o f S p e c i e s No. From i n t h e C y c l e s 3 l o s e Y i e l d l o s e Y i e l d P i t h I n c r e m e n t % % P s e u d o t s u g a m e n z i e s i i 5 0 0 - y e a r - o l d 80 EW 3 7-4 .8( 1 . 3 0 ) b 45. 2 ( 0 . 27) LW 5 74 .1 46. 3 P s e u d o t s u g a m a n z i e s i i NH .NO, f e r t i l i z e d 23 EW 3 74 .3 44. 9 LW 5 73 .9 45. 8 P s e u d o t s u g a m e n z i e s i i C o m p r e s s i o n wood 73 LW 5 61 .2 38. 4 S e q u o i a s e m p e r v i r e n s M a t u r e wood - EW 3 70 .1 43. 3 LW 5 68 .9 44. 2 P i c e a s i t c h e n s i s M a t u r e wood 36 EW ' 3 74 .0 46. 0 . LW 5 73 .4 47. 5 ( 0 . 29) P i n u s l a m b e r t i a n a M a t u r e wood — LW 5 72 . 2 ( 1 . 53) 46. 4 a E a c h c y c l e c o n s i s t s o f a 30 m i n u t e t r e a t m e n t w i t h p e r a c e t i c a c i d , f o l l o w e d by a s o a k i n g i n h o t w a t e r (30 m i n u t e s ) b No. i n p a r e n t h e s i s i n d i c a t e s p e r c e n t a g e o f r e s i d u a l l i g n i n EW E a r l y w o o d LW Latewood 142. APPENDIX I I I MOISTURE CONTENT OF THE HOLOCELLULOSE AND ALPHA-CELLULOSE PULPS AFTER CONDITIONING IN THE CTH ROOM FOR OVER ONE MONTH E a r l y w o o d / H o l o c e l l u l o s e A l p h a - C e l l u l o s e S p e c i e s Latewood M o i s t u r e M o i s t u r e C o n t e n t Q. "5 C o n t e n t O. *6 A r a u c a r i a c u n n i n g h a m i i EW 10.66 10.15 LW 10.87 10.38 S e q u o i a s e m p e r v i r e n s EW 9.86 9.52 LW 9.79 9.62 P i n u s l a m b e r t i a n a EW 9.80 9.83 LW 10.04 10.12 P o d o c a r p u s d a c r y d i o d e s EW 9.74 9.47 LW 9.81 9.52 P i c e a s i t c h e n s i s EW 10.58 9.8 0 LW 10.73 9 . 54 Taxus b r e v i f o l i a EW 10.00 9.65 LW 10.03 9.84 J u n i p e r u s v i r g i n i a n a EW 9.80 9.60 LW 10.07 9.96 C e p h a l o t a x u s w i l s o n i a n a EW 10.20 9.31 LW 10.20 9.31 P s e u d o t s u g a m e n z i e s i i EW 10.73 9.75 I n c r e m e n t 10 LW 10.01 10.10 i n c r e m e n t 20 EW 10.08 10.11 LW 9.98 9.71 I n c r e m e n t 80 EW 9.99 10.07 LW 10.05 9. 94 I n c r e m e n t 150 EW 10.88 10.16 LW 10.98 10.24 I n c r e m e n t 3 00 EW 9.99 9.90 LW 10.01 9.73 I n c r e m e n t 400 EW 10.59 9.92 2 LW 10.11 9.96 P s e u d o t s u g a m e n z i e s i i EW 10.26 9.52 3 LW 10.28 9.54 P s e u d o t s u g a m e n z i e s i i EW 10.34 9.43 I n c r e m e n t 20 LW 10.35 9.46 I n c r e m e n t 21 EW 10.31 9.61 LW 10.34 9.64 I n c r e m e n t 22 EW 10.19 9.40 LW 10.18 9.46 ( C o n t i n u e d ) 143. APPENDIX I I I ( C o n t i n u e d ) E a r l y w o o d / H o l o c e l l u l o s e A l p h a - C e l l u l o s e S p e c i e s L a t e w o o d M o i s t u r e M o i s t u r e C o n t e n t Q, *5 C o n t e n t a *o I n c r e m e n t 23 EW 10.18 10.05 LW 10.28 9.86 I n c r e m e n t 24 EW 10.20 9.79 LW 10.18 9.78 I n c r e m e n t 25 EW 10.19 9.88 LW 10.88 9.90 P s e u d o t s u g a m e n z i e s i i EW 9.85 9.49 I n c r e m e n t 15 LW 9. 86 9.52 I n c r e m e n t 16 EW 9.88 9.53 LW 9.85 9.52 P s e u d o t s u g a m e n z i e s i i EW 10.47 9.56 I n c r e m e n t 15 LW 10.50 9.58 I n c r e m e n t 16 EW 10.42 9.50 LW 10.48 9.52 P s e u d o t s u g a m e n z i e s i i EW 10.62 9.64 I n c r e m e n t 15 LW 10.59 9.62 I n c r e m e n t 16 EW 10.58 9.59 LW 10.57 9.61 1. B u t t l o g o f a 500-y e a r - o l d t r e e 2. C o m p r e s s i o n wood - B r e a s t '. h e i g h t d i s k o f a 1 2 0 - y e a r - o l d t r e e 3. NH.NO_ f e r t i l i z e d - B r e a s t h e i g h t d i s k o f a 2 9 - y e a r - o l d t r e e ; 4 j f e r t i l i z e d a t ages 21 and 22 (500 l b / a c r e 4. U r e a f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t , 2 4 - y e a r - o l d t r e e ; f e r t i l i z e d a t ages 13 and 15 (200 l b / a c r e o f N,P 20,.orK 0 e q u i v a l e n t s ) 5. NPK f e r t i l i z e d - 12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t , 2 4 - y e a r - o l d t r e e ; f e r t i l i z e d a t ages 13 and 15 (200 l b / a c r e o f - N , P 2 0 5 o r K 2 0 e q u i v a l e n t s ) 6. ( N H ^ ) 2 S O ^ f e r t i l i z e d -.12 mm i n c r e m e n t c o r e f r o m b r e a s t h e i g h t . 2 4 - y e a r - o l d t r e e ; f e r t i l i z e d a t .ages 13 a n d 15 (200 l b / a c r e o f N,P 20 o r K 2 0 e q u i v a l e n t s )

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