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Weight-length relationships of coniferous wood tracheid skeletons Sastry, Cherla Bhaskararama 1971

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WEIGHT-LENGTH  R E L A T I O N S H I P S OF  CONIFEROUS  WOOD T R A C H E I D S K E L E T O N S  by  C H E R L A BHASKARA M. M.  S c . (Botany)' Andhra  Sc. (Forestry)  A  RAMA  THESIS THE  University  SASTRY  University,  of British  SUBMITTED IN PARTIAL REQUIREMENTS  1958  Columbia,  F U L F I L M E N T OF  FOR T H E D E G R E E  OF  DOCTOR OF P H I L O S O P H Y  in  the Department of Forestry  We  accept  required  THE  this  thesis  as c o n f o r m i n g  to the  standard  UNIVERSITY  OF  April,  B R I T I S H COLUMBIA 1971  1967  In p r e s e n t i n g t h i s  thesis  an advanced degree at the L i b r a r y s h a l l I  f u r t h e r agree  in p a r t i a l  fulfilment of  the U n i v e r s i t y of B r i t i s h  make i t  freely available  that permission  for  the requirements f o r  Columbia,  I agree  r e f e r e n c e and  f o r e x t e n s i v e copying o f  this  that  study. thesis  f o r s c h o l a r l y purposes may be g r a n t e d by the Head of my Department o r by h i s of  this  written  representatives. thesis  It  for financial  i s understood that gain s h a l l  permission.  Department  of  The U n i v e r s i t y o f B r i t i s h Vancouver 8, Canada  Columbia  not  copying o r p u b l i c a t i o n  be allowed without my  ABSTRACT The tracheids  hypothesis  of coniferous  alpha-cellulose their  length,  earlywood  and latewood  skeletons.  skeletons  developed  with  750 i n d i v i d u a l  ultra  curvilinear of  NaBH^.  holo-  micro-balance,  to weigh; individual Statistical  separated  having  A  to  trac-  specially range o f  ± 0.03 y g , w a s c o n s t r u c t e d  and  tracheids.  analyses  indicated a significant  fraction.  positive  o f t r a c h e i d and w e i g h t  Estimated  variations  length  accounted  weights,  by  factor.; a l o n e w e r e , r e s p e c t i v e l y , 91.9 a n d 95.7 p e r  f o rpooled  data  of a l l the species.  i n holocellulose skeleton  skeletons  alpha-cellulose  within species,  between s p e c i e s ,  No s i g n i f i c a n t  weights were e v i d e n t  f o r t h e same t r a c h e i d l e n g t h ,  cellulose  peracetic  a weighing  the  species  into  reduction  and w e i g h t .  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  ferences  with  Further  for  cent  wood  and a l p h a - c e l l u l o s e  r e l a t i o n s h i p between l e g n t h  i t s carbohydrate  representing  families,  f o r length  0.06 t o 14 y g a n d a p r e c i s i o n used  o f h o l o c e l l u l o s e and  f o ra portion of the holocullulose  were measured  quartz  individual  f r a c t i o n s , was d e l i g n i f i e d  followed  About  increments,  genera and seven  and subsequent r e d u c t i o n  alpha-cellulose  the weights  among  species.  o f 28 a n n u a l  coniferous  that,  are a direct quantitative function of  independent of  from nine  heid  woods,  skeletons  A total  acid  i s examined  whereas weights  and both  differed  the holo-  difwithin  of  alpha-  and  significantly.  Radial trends  similar  gravity,  variation  t o those  and p e r c e n t a g e  for single  established  Individual  tracheids  lower  (1% l e v e l )  alpha-cellulose  mature  significant  lighter  m a t u r e wood,  Differences For  t h e same  heids  tracheid  Examination  weights of  (normal)  differences that,  were  non-  tracheids  weight)  than  were those  and latewood  were e x p l o r e d .  and latewood  trac-  was  higher  wood  also  wood  wood,  tracheids  provided positive  i n tracheids.  were compared  no s i g n i f i c a n t tracheid  i n latewood.  differences  skeletons,  length,  signifi-  I t was  compression  lower  amount o f a l p h a - c e l l u l o s e  wood,  b u t a h i g h e r amount t h a n  those  were  whereas  f o r alpha-cellulose.  tracheid  with  When  con-  wood  than those  those from  wood.  Changes wood  those  length.  earlywood  per tracheid  f o r t h e same  (normal)  than  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  were found  may.have a  mature  juvenile  in  both  for holocellulose  tracheids from  length,  of compression  of compression  apparent  cluded  wood  f o r the length-weight r e l a t i o n s h i p  regular  cant  tracheid  earlywood  of holocellulose  evidence  weights  Overmature  analy-  significantly  differences  between  contained similar  amount  skeleton  had  (carbohydrate skeleton  f o r t h e same  followed  on g r o s s wood  wood  wood, w h i l e  forholocellulose.  significantly from  of juvenile  and overmature  weights  by o t h e r s f o r s p e c i f i c  of c e l l u l o s e based  ses.  from  tracheid  formed  i n Douglas-fir  tracheid  b e f o r e and f o l l o w i n g  tree  weights  were  fertilization.  studied  Variations,  f o r t h e most  changes  i n tracheid  butable  to treatment  were of  also  noted,  with  n o r m a l wood  tion  was  in  gross  length This  gravity  differences  reduction factor  with  vs.  resulted  of tracheid,  i n wood  compatible  t o be a s s o c i a t e d  ( U r e a v s . NPK  treatments  as a p o s s i b l e  were  found  Qualitative  some  per unit  specific  Results  were  composition  tracheids.  suggested wood  length.  i n that  alpha-cellulose  part,  attri(NH^^SO^)  i n less  when  for differences  fracobserved  fertilized  the proposed  weight  compared  i n cellulose  o f some  with  trees.  hypothesis.  iv  TABLE  OF  CONTENTS Page  ABSTRACT TABLE LIST  OF OF  LIST,OF LIST  i CONTENTS  i v vi  TABLES FIGURES  v i i i  OF A P P E N D I C E S  x  ACKNOWLEDGEMENT  x i  CHAPTER I. II.  INTRODUCTION  1  L I T E R A T U R E REVIEW  4  Tracheid  Morphology  Tracheid  Length  Dimensional  4  Variation  11  Relationships  i n Tracheids  Chemical Composition and Recovery Carbohydrate F r a c t i o n Estimates of Carbohydrate in Relation to Cellular Effects and III.  of F e r t i l i z a t i o n  Chemical  MATERIALS  AND  Selection Wood  19  26  Properties  Composition  29  METHODS  35  of Material  35  Sample  Analysis  of  F r a c t i o n and Weight Elements o n Wood  15  and T r a c h e i d  of Tracheid  Skeleton  Skeletons  Preparation  37 40  1.  Sampling  criterion  40  2.  Tracheid  length  41  3.  Tracheid  weight  42  V  CHAPTER  Page Statistical  IV.  RESULTS  AND  Analyses  45  DISCUSSION  47  Relationship  Between Tracheid  1.  Variation within  2.  Variation  3.  Variation with  4.  Variation and  Some P r a c t i c a l  VI.  species  between species increasing  52 age  between r e g u l a r  55  (normal)  between earlywood  of the Relationship Considerations  RECOMMENDATIONS FOR  48 48  f o r a Common R e l a t i o n s h i p  Applicability  V.  and Weight  c o m p r e s s i o n wood  5. . V a r i a t i o n Evidence  Length  FURTHER RESEARCH  60 and-latewood  64  i n Conifers  68 72 82 86  CONCLUSION  88  LITERATURE CITED  91  TABLES  105  FOREWORD TO F I G U R E S  120  FIGURES  121  APPENDICES  139  vi L I S T OF T A B L E S NUMBER I. II.  III. IV. V.  VI.  VII.  VIII.  IX.  X.  XI.  Page Single tracheid (holocellulose skeleton) 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  weights 105  Single tracheid (alpha-cellulose skeleton) 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 Single tracheid (holocellulose skeleton) 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  weights 107  weights  Single tracheid (alpha-cellulose skeleton) and l e n g t h s f o r t w o D o u g l a s - f i r t r e e s  109 weights 110  A n a l y s i s m o d e l (A) a n d (B) r e s u l t s o f m u l t i p l e curvilinear covariance analyses; differences among s p e c i e s (holocellulose skeletons). (Adjusted f o rdifferences i n tracheid length)  111  A n a l y s i s m o d e l (A) a n d (B) r e s u l t s o f m u l t i p l e curvilinear covariance analyses; differences among s p e c i e s (alpha-cellulose skeletons). (Adjusted f o rdifferences i n tracheid length)  112  Multiple curvilinear covariance analysis f o r tracheid weights (holocellulose skeletons) 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 500^year-old Douglas-fir tree.. (Adjusted for differences i n tracheid length)  113  Multiple curvilinear covariance analysis f o r tracheid weights (alpha-cellulose skeletons) 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 500-year-old Douglas-fir tree. (Adjusted for differences i n tracheid length)  114  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 of earlywood and latewood t r a c h e i d weights i n six Douglas-fir trees. (Adjusted f o r d i f ferences i n tracheid length)  115  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 of earlywood and latewood t r a c h e i d w e i g h t s i n nine coniferous species. (Adjusted f o r d i f ferences i n tracheid length)  116  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 e r t i l i z e d Douglas-fir trees  117  of three  vii  NUMBER  XII.  Page  R e s u l t s of 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 analyses; differences within species in t r e a t e d and u n t r e a t e d Douglas-fir. (Adjusted for differences i n tracheid length)  118  v i i i L I S T OF  FIGURES  NUMBER 1.  2.  3.  4.  5.  6.  7.  8.  9.  Page R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 samples, from Araucaria cunninghamii  121  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 Sequoia sempervirens  122  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 ) a n d l e n g t h f o r samples^" f r o m Pinus lambertiana .  123  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 (holoc 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 ) a n d l e n g t h f o r samples"'.from Podocarpus dacrydioides  124  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 ) a n d l e n g t h f o r samples.: f r o m Picea sitchensis  125  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 f r o m Taxus b r e v i f o l i a ,  126  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 ) a n d l e n g t h f o r samples-" f r o m Juniperus virginiana  127  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 samples." f r o m Cephalotaxus wilsoniana  128  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 samples from Pseudotsuga menziesii  129  ix NUMBER 10.  11.  12.  13.  14.  15.  16.  17.  18.  Page R e l a t i o n s h i p between t r a c h e i d weight (holoc 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) skeletons and - length f o r samples from nine coniferous species. (Least-squares fitted line)  130  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 samples from nine coniferous species. (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 pass through t h e origin)  131  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 , across t h e b u t t l o g o f a 500year-old Douglas-fir tree. (Each p o i n t r e p r e sents the average t r a c h e i d weight o r length f o r the increment. S p e c i f i c g r a v i t y - values from Kennedy and Warren (70))  132  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 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  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 length patterns across s i xconsecutive increm 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 c o m p r e s s i o n wood 4 y e a r s a f t e r first fertilization. (Each p o i n t represents the average t r a c h e i d weight o r length f o r t h e increment)  134  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 Urea t r e a t e d and normal D o u g l a s - f i r  135  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 ) a n d l e n g t h i n NPK t r e a t e d a n d normal D o u g l a s - f i r  136  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 (NB^^SO^ 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  R e l a t i o n s h i p between t r a c h e i d weight (holoc 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 ) a n d l e n g t h i n NH4NO3 t r e a t e d a n d normal D o u g l a s - f i r  13 8  X  L I S T OF A P P E N D I C E S NUMBER I. II.  III.  Page Description the study  o f growth  increments  included i n 13 9  Summary o f 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 y i e l d s a n d 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 samples s t u d i e d (based on ovendry, e x t r a c t i v e f r e e wood)  141  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 and a l p h a c e l l u l o s e pulps a f t e r conditioning i n the CTH Room f o r o v e r o n e m o n t h  142  xi ACKNOWLEDGEMENT The  writer  acknowledges  the continuous  inspiration  and c o n s t r u c t i v e c r i t i c i s m  entire  at the University  stay  supervisor, him  The ledged,  his  help  and kindness  Columbia,  Without  phases  or  suggestions  of the thesis,  enlightening inspiration,  by h i s  the interest  shown by  completed.  o f D r . J.W. W i l s o n  f o r h i s many u s e f u l  and experimental  truly  to him during h i s  might n o t have been undertaken  not only  planning  of British  D r . R. W. W e l l w o o d .  the thesis  given  guidance,  i s acknow-  during the but alsof o r  encouragement and p e r s o n a l  concern. Dr. questions gratefully  concerning  thewriter  statistical  i nclarifying  analyses.  many  H i s advice i s  acknowledged. The  committee  A. K o z a k h e l p e d  thesis  consisting  was r e v i e w e d  by members o f t h e a d v i s o r y  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 a n d D . J . W o r t . suggestions,comments  For their  and c o n s t r u c t i v e c r i t i c i s m  valuable  thewriter i s  indebted. Many h e l p f u l Washington U n i v e r s i t y Cabot Foundation  suggestions School  from  D r s . O.H.  of Medicine,  f o rB o t a n i c a l Research,  S t . L o u i s , D.S. Harvard  sham, B.S. Wenger, D e p a r t m e n t o f Anatomy, S c h o o l University  o f Saskatchewan,  and t h e time  Lowry, Skene,  Forest, Peterof Medicine,  and t a l e n t s  o f many  xii p e r s o n s on U.  Rumma, K.  ment o f the  the  the  Faculty  Apt,  and  quartz  G.  technical staff,  Bohnenkamp, made p o s s i b l e  ultra-microbalance,  an  integral  Messrs. developpart  understanding  their  advice  is  with  some o f  the  following contributed  the problems  involved herein  greatly, and  appreciated.  Dr.  R.W.  Dr.  L.  Paszner  Faculty British  of F o r e s t r y , Columbia.  University  of  Dr.  J. Worrall  Faculty British  of F o r e s t r y , Columbia.  University  of  Dr.  D.J.  Department of Botany, U n i v e r s i t y B r i t i s h Columbia.-  Kennedy  for  Hejjas  D e p a r t m e n t o f F i s h e r i e s and Canadian F o r e s t r y S e r v i c e Forest Products Laboratory Vancouver.  Wort  The K.  of  s tudy.  Discussions in  of F o r e s t r y  w r i t e r s i n c e r e l y t h a n k s M i s s L.  f o r w r i t i n g the  samples;  for technical assistance the N a t i o n a l  University.of  British  Finally, his wife,  Cho  he  and  i n the  Research C o u n c i l Columbia  C.P.  Chen,  e n c o u r a g e m e n t and  Mrs.  Lambden  Graduate  o f Canada and  of  pulp  the  assistance.  must e x p r e s s h i s s p e c i a l  Ratna, f o r her  M.  preparation  for financial  of  C o w d e l l and  computer p r o g r a m s ; Mrs.  d r a f t i n g work; M e s s r s H . J .  Students,  Forestry,  appreciation  monumental  to  patience.  CHAPTER  I  INTRODUCTION  Much emphasizes cellular which by  in  elements  weight  up  of coniferous  a s much  of such  understanding  a s 90 p e r c e n t  tion  within  radial  and  physical the  a single  mental  individual chemical  cell  as w e l l  Less  have  tracheids  o r 98 p e r  the tracheid  and study  cent  i s the  should  assist  activity  At s t i l l  Further level,  within  and between level.  studied  and  analyzed  another  has been  intensive  i n particular cell  including cell  walls.  spatial  zones  has  to  (corewood, earlywood  f o r differences i n of investigation,  properties  examined  study  respect  separate  level  and c h e m i c a l  species  Similarly, varia-  with  as wood  frequently,  been  increments  composition  constituents  has been  i n physical  growth  analyses.  Since  a t the gross  patterns,  behaviour.  individual  principal  are longitudinal  i t si s o l a t i o n  stem  segments  variability  The  by volume  i n wood w i t h i n  and sapwood).  latewood  level.  science  o f wood.  extensively  and h e i g h t  heartwood  material.  woods,  Variability studied  woods  and pulp  and i n t e r p r e t i n g the b i o l o g i c a l  behaviour  been  i n wood  a t the c e l l u l a r  of the ovendry  unit  product  research  variability  make  basic  current  by  intra-incre-  occurred wall  within  at the  organization  distribution  of the  and  2.  Even ties is  o f wood  known  vidual  though  have  about  been  characteristic,for has been  coarseness,  or tissue  weakness i n t h a t  industries  deduced  efforts  expression  have  o f wood  important  related  lead  indicators, Also, fibre  1  on p u l p  been  knowledge  in  order  methods  each  changeably  Though  such  such  have  segments  indi-  measured  as  an  fibre  inherent  variation  o f t h e wood  wood  quality.  f o r an a n a t o m i c a l  i s generally  basis  of cellulose  content  by v a r i o u s  amount, w i t h i n  for  on t e n s i l e  as  length  of  individual  established. strength a  and u n d e r s t a n d  The  o f wood thorough  of fibre  species,  'tracheid'  quality  properties.  Certainly,  and between  and  strength  content  closely  are used  has been  between  t o be  and s t r e n g t h  workers.  evaluate  "*"The w o r d ' f i b r e ' i n this thesis.  that  length  properties  carbohydrate  known  and v a r i o u s  of tracheid  fully  and t h e i r  characteristics  and paper  t o more  per  cellular  of defining  to a search  of the relationship  carbohydrate  methods  i n various  as d e n s i t y ,  confirmed  length.  of tracheids  wood  the influence  significance has  such  example,  little  quality.  because  to gross  t o an e a s i l y  these  proper-  relatively  fraction  of individual  Scientists  Dimensions are  studied,  relation  fractions,  are emphasizing  and p h y s i c a l  by i n d i r e c t methods  the estimate  not possible.  These  intensively  and p o s s i b l e  information  is  constituents  t h e amount o f c a r b o h y d r a t e  tracheid,  anatomical  chemical  and i t s  i s necessary  gross  are used  wood  inter-  properties  and t o more f u l l y  ledge regarding exponential),  utilize  wood.  the type of r e l a t i o n s h i p  w o u l d be o f i m p o r t a n c e  In a d d i t i o n ,  (i.e.,  linear or  t o g e n e t i c i s t s who a r e  e n d e a v o u r i n g t o m a n i p u l a t e wood c h a r a c t e r i s t i c s t h r o u g h b r e e d i n g work.  E v e n more i m p o r t a n t ,  m e n t a l knowledge s h o u l d  know-  any g a i n s  be o f g e n e r a l  value  i n this  tree funda-  t o wood-based  technologies.  The 1.  objectives  To t e s t t h e h y p o t h e s i s  thesis are: that,  among i n d i v i d u a l  tracheids  of coniferous  cullulose  and a l p h a - c e l l u l o s e s k e l e t o n s  quantitative of 2.  of this  a.  woods, t h e w e i g h t s  function of their  length,  of holo-  are a d i r e c t independent  species. To a p p l y  the r e l a t i o n s h i p within  Douglas-fir by  a single  (Pseudotsuga m e n z i e s i i  comparing  regular  (Mirb.)  (normal) wood w i t h  species, Franco)  juvenile  wood, o v e r m a t u r e wood, c o m p r e s s i o n wood, and wood formed w i t h i n  trees  treated with  chemical  fer-  tilizers . b.  To examine t h e r e l a t i o n s h i p w i t h i n by  increments  comparing i n d i v i d u a l carbohydrate  from earlywood  and l a t e w o o d .  skeletons  CHAPTER I I  LITERATURE  During anatomical and  structure,  physical  increasing subject  the past  been  127,  136, 153, 1 6 8 ) .  supplemented  same g r o u n d .  An  of  which  Tracheid  introduced  accumulated  have  composition,  i n ever-  been w r i t t e n on t h e  I t would will  papers  seem  are related  these  ( 1 9 , 20, 25, 31, 34, 70,  superfluous  b e made  to cover  directly  stated  by Wardrop  originally  that  field  (145),  by G o e t h e  to re-cover the only  those  to the thesis  of study  concerned  as d i s t i n c t  from  scope  o f the term  has been widened  structure  and form,  structure  even  concerned  the study  and more  points  topic.  also  with  internal  with  the physiology  wood  structure o f wood  with  plant  of function.  century  and With  time,  fibre  the both of  discussion  e x t e r n a l geometry b u t  appropriate)  formation.  to  of  to i n c l u d e the study Therefore,  was  animal  t o i n c l u d e the study  level.  (where  morphology  i n the l a s t  recently  a t the molecular  not only with  the term  early  form,  is  chemical  ( 4 5 , 62, 72, 1 0 1 , 137) a n d  by review  attempt  regarding the  Morphology  As  define  have  S e v e r a l books  have  data  properties,  o f wood  within recent years  literature  t o 50 y e a r s  physical  functioning  volume.  40  REVIEW  and, b r i e f l y  5. The the a  vascular  p r o d u c t i o n o f wood r e s u l t s cambium w h i c h  s h o r t d i s t a n c e below  branches  of a tree  vascular  cambium:  Fusiform the  fusiform  2)  ray  shape  (19).  The  elongate  elements  initials  produce  of  the xylem  Two  and  a r e much s m a l l e r former of  the  a l l elements phloem  and of  and  Periclinal about  initials  the xylem  of  is particularly vascular  and  the  i n the  almost  the  ends,whereas  isodiametric  to a l l the  phloem, whereas  in  vertical  the  transverse or ray  division  of  or  occur  radial  add  ray system  w h i l e the  cambial c e l l s expansion  their  and  initial  affecting quality  important to conifers  elements  anticlinal  i s traceable  back  and  or  and  latter  with  (8).  to a l l the v e r t i c a l  factors  to the kind  and  vascular  to the xylem  growth,  fusiform  phloem,  i n the  additive  divisions  give rise  and  great importance  derived  types  tapering  give rise  and  periclinal  t h e many p h y s i o l o g i c a l  This  cell  with  accommodation t o c i r c u m f e r e n t i a l  of  t h e main stem  (45).  in multiplication  fusiform  ments  initials  the xylem  phloem hence bringir.  quent  continuous  and  are elongated c e l l s  cambium o f c o n i f e r s ;  result  a r e two  initials;  types of c e l l  multiplicative.  t i p of  of  initials.  initials  ray i n i t i a l s  There  activities  circumferentially  the growing  (45).  1)  becomes  from  type  conseBecause  elongate  final  ele-  dimensions,  the dimensions, are o f wood  since  the  produced. length  to cambial  of  initials  6.  from which gation  of  tracheids daughter  were  cells  derived,  i . e . , post-divisional elon-  is relatively  slight  in  conifers  ( 2 , .8) .  The has  been  extensively  Wardrop, this  in  area  two  (143,  synthesis his  pattern  of  studied  involves  The  144,  four  bium,  10  to  20  cell  2)  enlargement  division  begins, are  the  4)  lignification  complete, then  as  or  by  from  fibres  the  follows  work  and  of  work is  a  in brief  Wardrop  and  148).  the  cambium  of  conifer  daughter  of  cells  to  the  dimen-  cell; the.secondary  result  of  to  500  cell  per  wall;  the  toward  middle their  secondary  division  dimensional cent  Development  dimensional  near  the  cambium;  considerable  200  before  of  in  and  (147).  more.  extends  development  a  undergo  first  that  147,  the  the  development  radially cent  146,  mature  the  i n part,  tracheids sive  of  in  of  3)  per  summarized  extensive  tracheids  1)  immediately  formation  phases:  c e l l s , formed  increasing  and 145,  of  has  discussion  excellent  (143,  wall  A u s t r a l i a n workers,  144).  sions  Daughter  by  articles,  Development stems  secondary  recent  the  associates  of  of  changes of  the  tips.  and  in  in  cam-  change, length  secondary of  the  by  wall  differentiation  differentiating During  wall, contact  the  between  progresthe  7.  primary  wall  terminal Wardrop cell in  canal and  wall  the  and  the  between  Dadswell  i n mature  middle.  the  cell  t i p , also  per  se  length  The cell are is  various  enlargement s t i l l  a  not  and  fully  function  of  that  this  is  time  that  cell  was  the  wall  secondary the  to  tracheid  body.  factors  which  the  enlargement  from  Cell  water  a  and  wall  the  the  (147).  that  similar  than  the  pro-  i t has  towards  the  tips  cell  been  of  the  (14).  As  noted  of  extent  enlargement  the  tips  portion  the  following  of  middle  regime  occurs  minute  supporting  influence  development  vacuolation to  at  possibility  compared  a  thickening  less  of  understood.  related  that  often  by  primary  besides  wall  cell  the  observation,  indicates  the  and  reported  weigh : l e s s  from  i s maintained  lumen  (146)  This  development  should  the  tracheids  gressive  unit  protoplast  of  division  apparently suggested  trees by  at  the  Wardrop  (148) :  In the l i v i n g c o n d i t i o n the c e l l s o f the cambium e x i s t i n a s t a t e o f t u r g o r and the 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 det e r m i n e d by t h e d i f f e r e n c e b e t w e e n 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 (potential) pressure a v a i l a b l e through osmotic a c t i v i t y of substances i n the c e l l sap.  Cell intrusive one  or  101).  growth,  both  ends,  According  tracheid  elongation  of  fusiform  i.e., tips between to  the  current  i s regulated  by  of  is  achieved  the  elongating  cell  other  developing  cells  concepts,  auxin  initials  radial  emanating  from  extend (2,  expansion growth  of  by at  9,  45, the  centers  8.  in  t h e crown  the  multinet  theory  During slip  theory  implies  bundles  by  (77, 8 0 ) .  cell  along  expansion, each  subsequently  orientation  and  The  down  (145). ferences strands  S^r This  wall  i s moved  structure.was  developed  wall  S^--of  into  heterogeneity  the mature  concept  and wall  thickening  size  layers—an  and  shape).  outer  optical  has been  wall  and  by K e r r  (secondary  differing  transverse  the primary  i t sfinal  of three  become  the primary  accepted  wall  outward  expansion.  The p r e s e n t l y  the c e l l  and  However, t h e  f o r the  wall  structures:  of  layer,  properties  attributed  to d i f - .  orientation of the c e l l u l o s e micellar  i n the three  innermost  displaced  o f i t sdevelopment,  has reached  and an i n n e r  i n helical  consists  and t h e s t r a n d s  to cell  o f two  consists  optical  sheath  This  microfibrils.  are passively  an e x p l a n a t i o n  prior  divided  the c e l l  wall  oriented  material,  and the secondary  after  secondary  middle  the  wall)  of the wall  membrane  not offer  wall.  They  part  formulated  wall.  i n the d i r e c t i o n o f growth.  consists  of c e l l  (67).  (cambial  wall  the standpoint  the secondary  Bailey  as t h i s  (115)  of the primary  the f i b r i l s  of the f i b r i l s  tracheid  terminology  laid  does  Roelofsen  transversely  o f new  oriented  From  a  other  theory  conifer  the innermost  or less  the apposition  multinet  f o r the growth  that  o f more  In addition,  layer  layers may  excrescences  (wart  protoplasmic  debris.  (143, 144>r145).  be s c u l p t u r e d  structures) These  In conifers,  i n the form  (85, 1 4 5 ) , p o s s i b l y  aspects  have  been  of  small  formed  extensively  from  9.  reviewed model (36)  by W a r d r o p  of the c e l l recently  electron  that,  From  other  thickest  the  volume  studies  workers  f o r both  or weight  in  of the  sion in  wood  their  They  some  tracheids typical  usually  fibrillar  wood  between  and  wood  lacks  often  layers  contains  highly to  next  little  f o r 80  (63) a n d  i t i s understood  t h e S2  layer.is  by.far  t o 95 p e r c e n t  i n the wall  of  organization,  as i n  Compres-  i n cross  appearance layer as w e l l wood  contain  layer  section  normal  and i n t e r c e l l u l a r and numerous a s a much  an e x t r a  of the three  radial  micro-  of l i g n i n (13 6 ) ,  located  wall  layer,  gelatinous  and c o n s i s t s  com-  tension  secondary  an a d d i t i o n a l w a l l  i f any l i g n i n  spaces.  In addition,  to Timell  even  ones  regular  greater  layer  the so-called  microfibrils  (53) .  from  tracheids.  According  possesses  a x i s "(136).  them,  i n particular  an  t o t h e lumen,  crystalline  the f i b r e  t o by  one o r s e v e r a l  and f r e q u e n t l y  deposited This  (136).  and F e n g e l  provided  spp. l a c k  normal  tracheids  and  that  t h e S^r  than  fibres,  and  and i t i s r e p o r t e d  h a v e ,a w i d e r  angle  Dunning  anatomy,  differ  rounded  discontinuities.in  pression  Picea  schematic  fibre.  do o c c u r  o f r e a c t i o n wood  n o r m a l wood  o f Jayme  accounts  a  In addition,  organization,  and latewood,  and p r o b a b l y  developed  f o r microdissection  pulp  wall  referred  earlywood  Variations case  a.new m e t h o d  on c e l l  who  144).  o f s i n g l e wood  information  the  the  (142, 143,  developed  layer.  several  wall  microscopy  additional the  and h i s a s s o c i a t e s  layer.  largely of  of cellulose, oriented  S i m i l a r l y , the morphology  of  parallel tracheids,  10.  i.e.,  dimensions  earlywood There wall  and  and  thickness of wall  latewood  o f t h e same g r o w t h  i s an i n d i c a t i o n , area of earlywood  constant  i n conifers  According Stewart  (127),  classified  however,  (14,  differs  increment  between  (137).  that the cross-sectional  and o f latewood  t r a c h e i d s may  cell  remain  68).  t o Wardrop  the c e l l  wall  three  groups:  into  layers,  (143)  chemical  and as s u m m a r i z e d  by  c o n s t i t u e n t s may  be  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 n d 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 ) polysaccharides are deposited b e f o r e t h e p r i m a r y ( l i g n i n ) and secondary (extractives) encrusting substances. The  distribution  uneven  throughout  lation  of the middle  and  indicated  the c e l l  wall.  lamella  Bailey  constituents i s  (1) r e p o r t e d t h e  Lange  and h i s associate.'.  s p e c t r o p h o t o m e t r i c method Swedish  spruce  (Picea  earlier  findings  (L.) K a r s t ) ,  of Bailey.  16 p e r c e n t  reviewed  this  aspect  compound m i d d l e  and  (75)  and  applied  and c o n f i r m e d  In addition,  concentration external with  lignin,  14 a  per  micro-  on 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  abies  compared  iso-  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  t h a t i t c o n t a i n e d 71 p e r c e n t  cent pentosans.  lignin  of the chemical  near  to the  layer  the lumen.  concluded  i t was  noted  the that the  i s 73 p e r  cent,  B e r l y n and Mark  (11)  t h a t although most of the  lamella, i s ligniri;-.this  mean t h a t m o s t - o f t h e - l i g n i n ' (amount). .is :  does  not n e c e s s a r i l y-  i n this  region.  11.  Information saccharides individual deduced (90,  recent this  i n the c e l l  the spatial wall  polysaccharides  very  91)  on  roughly  by  d i s t r i b u t i o n of  is limited.  i n each  following  cell  area  as  Panshin  and-de  wall  proportion layer  microdissection  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  book,  The  Zeeuw  methods  (101)  poly-  had  of  been  techniques  (74).  summarized  In  their  t h e work  in  follows:  . . . 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 is c e l l u l o s i c ; i n the S layer the c e l l u l o s e i n c r e a s e s t o m o r e t h a n 50 p e r c e n t o f t h e material i n that c e l l wall layer; a reduction 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^. ?  It major  component  cellulosic wall  i s clear  those  of the l i g n i n  lated  inversely  Tracheid  effects  f a r has  components little  components  per  Length  that  c e l l u l o s e i s the  whereas  dominant  that  has  ' i n the  mostly  on  the  non-  primary with  of l i g n i n  polysaccharides  i n the tracheid,  attempt  and  of these.values  percentage  centred  lignin  is re(101).  spatial  i.e., qualitative  b e e n made  to quantify  the  tracheid.  Variation  extensive  variations of  wall,  of the t o t a l  but very  An length  so  studies  a comparison  indicate  to that  done  distribution.of  various  are  Furthermore,  T h u s .the w o r k  these  i n the secondary  polysaccharides  (145).  aspects,  from  both  the external  literature within  has  accumulated  and between  environment.  The  trees  on  tracheid  and on  pioneering  probable  work  i n  this  area  tracheid Within come  i s traceable length  quent  century  expansion.of As  31, 1 2 2 ) .  able  on t h e p a t t e r n s the process  some  a result,  r e l a t i o n s h i p s a r e now  radially serial  within  sections  ception  between  that  of yearly  first  cell  a growth  growth  portion  increases  Thus,  latewood  longer  than  vertically  earlywood confirmed tracheids  earlywood within  gradual  increase  crown,  o r between  upon  decreases  following  i s reached  the nature zones  that  of  follows upward  3 0 to 4 0 per cent  macerating the i n either i n  transition  from cited  about  a general  studies,t h e same (31, 1 2 2 ) .  10 p e r  length  cent  variation pattern  t o a maximum  of the tree  the  either  Other  material  Tracheid  evident  Thereafter,  (15).  a r e on t h e a v e r a g e  the base  tree  t o the boundary  the r e s u l t s already  tracheids.  a  so  are  shown by  or close  and latewood  an increment from  changes  i n the latewood,  t h e two i n t r a - i n c r e m e n t a l  have  length  t o a maximum  depending  i s avail-  zone  (15, 3 1 ) .  between  increment,  and hardwoods  and latewood  or slowly,  isolated  conse-  a  variation within  a minimum  rapidly  using  with  information  I t has been  length until  has be-  clear.  of the earlywood  the earlywood  length  cell  length  i n the cambial  of cell  ring.  (4) o f  s y I v e s t r i sL . ) .  conifers  length  quite  patterns  industry,  sufficient  division  (Pinus  of fibre  on b o t h  of tracheid  of c e l l  Definite  the  and paper  information  investigations  pine  the matter  to the pulp  (30,  and  first  v a r i a t i o n i n Scotch  the present  important  to Sanio's  of  below t h e  height  (9,,30  13. 31,  82, .98) a n d a d e c r e a s e  the  trunk. Radial  growth  increments,  studies, given very  height,  1.5 mm  increase  coniferous  may b e r e a c h e d  of  reached  length  considerably  both parts length  Furthermore,  t o bark,  from  to five  (137) .  throughout  growth  length i s times  maximum Maximum  greater  t h e growth  (46) .  finally,  continuincrements  vary  i n very o l d  t h e maximum  but earlywood  the variability  length  l e n g t h may  t r a c h e i d s have a g e n e r a l  than  length  A f t e r maximum i s  length; from  r u l e as  a n d may n o t  o f age; a  during which  that  0.5 t o  the rate of  as s i x t o e i g h t y e a r s ,  i n the.early years  i s much l e s s t h a n  increment are  a maximum  i s 200 t o 300 y e a r s  of the increment,  (155).  then  f o rreaching  l e n g t h may d e c r e a s e  rise  cells  necessary  follows a period  from p i t h  growth  A t any  T h e r e i s no g e n e r a l  Douglas-^fir tree  gradual  wood  until  a b o u t a mean maximum  the c e l l  number o f  r a p i d l y i n t h e second  (26, 3 1 ) .  has been observed  there  i n successive  range i n length  the r a t e of growth  a tree  a 455-ryear-old  trees,  first  may b e t h r e e  i n as l i t t l e  until  increase  reached,  in  increases  t h e number o f y e a r s  ous  of-the  tracheids  In conifers this  varies with  height i n  a r e summarized below.  declines, but continues  the i n i t i a l  which  be  of which  and i n a few subsequent ones;  reached.  to  of tracheid length,  the tracheids  (31). Length  increment  above t h i s  has been i n v e s t i g a t e d i n a g r e a t  the results  short;  than  variation  i n length  (5). In increase  e x h i b i t s a more  i s shown by t h e l a t e w o o d i n length  f o r latewood  f o rthe early- . cells  (30,  155).  14.  There relationship of  opinion  within such  a  as  and  being  length two  height  32,  82,  of  during  the  was  has  to  i s also  113),  control  was.  negative  shorter  122, found  to  from  growth  to  have  the balance  i.e.,  other  factors  regards  height  between  pith;  rates,  the  one;  As  exist  to the  cells  137).  increment  shown  of  longer  this  thus .in  the  variation  tree  of  more  tracheids  which  modified of  by  be  that  evidence (8,  tree,tracheid length Dinwoodie  within  the  strongly  i n f l u e n c e d by  probably  by  means  of  (32)  tree the  is  by  (7,  been  37,  47,  a  and  slight  function of of  the  from  forma-  place  cell  length  (8).  How-  modifications agents  presented  "that  modi-  particular  100,  c o n t r o l l e d by  concluded  the  takes  location  61,  be  direction  environmental  has  is  as  in a  although  activity  auxin."  trees  to  involve  (145),  geographical  induced  length  such  wood  between  emphasized can  tracheid,  compression  length  tracheid  variation  found  changes  be  Thus  often  environmental  influenced  factors.  are  i n growth  considerable  the  40,  but  changes  length  over  a  length,  undergone  Tracheid  to  of  respect  106).  formation  i t is  Within  (31,  cell  have  different  patterns  due  tracheid  112,  favor  first  of  structurally  species  of  in  to  increments  the  growth  tree  decreases.  ever,  in  86,  i f the  vertical, tion  be  equal  trees  The fied  to  agreement with  width  a positive relation  comparing  23,  general  increment  tree, wider  cell  rapid  of  appears  age  growth,  i s no  for  (108,  genetic  154,  169).  physiological  tracheid  cambium terminal  length  which  is  meristem,  Studies in  length  on t h e mechanisms  w i t h i n the tree  i n v e s t i g a t i o n s o f Bannan anticlinal factor. of  be  Bannan, i n t h i s  ( 7 , 8,  i n variations  but the  extensive  9) h a v e s h o w n t h a t  appropriate  regard  result  are d i f f i c u l t ,  divisions taking place  I t would  that  i n t h e cambium t o quote  the rate i s a  a recent  of  principal  statement  (9):  In general, rate of a n t i c l i n a l d i v i s i o n and 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 . This 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 and t o species-to-species differences. However, some" notable exceptions occur. Thus i n S e q u o i a sempervirens, where t h e c e l l s a r e longer than in other conifers, the rate 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 above a v e r a g e . . Conv e r s e l y , Pinus c o n t o r t a has both r e l a t i v e l y short c e l l s and a low 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 unusual combination. It the  therefore cambium  which  cell  appears  that  i s probably  the rate of a n t i c l i n a l  a part  dimensions, determined  environmental  and o n t o g e n e t i c  Bannan's work  i s significant  of  cell  gations  of the operative  shape and dimensions of xylem  Dimensional  by  Relationships4Jn  diameter  mechanism  by  the interplay of  factors are regulated i n that with  i tbridges  genetic,  (9).  earlier  plant physiological  studies investi-  Tracheids  of v a r i a t i o n i n c e l l  t r e e were p r i m a r i l y concerned w i t h cell  i n  differentiation.  Investigations  of  division  and w a l l  These and r e l a t e d aspects  thickness  cell  size  length.  were g i v e n  on c o n i f e r s were  within the  Possible little  discussed  changes  attention. recently  by  Larson  southern wall  (77). pine  Goggans  species,  thickness  of cambial  ordinarily  very  Nevertheless, ably  tracheid  this  general  width  several  that  of diameter  and  determined  by t h e  are derived,  expansion  diameters  takes  place  increase  increase  and (2).  consider-  with  age on a  differences  rapid  authors  latewood  tends  since  from  the early  tracheid  t h e apex.  that,  This  formation  and t h e  and between  trees."  ( 9 5 ) was  the pith  to increase  years. with  diameter a t some  a single  usually  point  the  Larson  age,  applicable  increases  can be  i n mid-stem,  radial and  to earlyestablished  increment,  from then  the  (77) q u o t e d  increasing  growth  radial  and t h a t  t o a maximum w i d t h  i s primarily  Within  that  outward  no d e f i n i t e age p a t t e r n  tracheids.  t o a maximum  towards  during  within  and Roth  increases  d i r e c t i o n , and i t i s  t o latewood  i n t h e wood  o f Mohr  thereafter.  tracheids  i n the r a d i a l  and concluded  generally  earlywood  occurs  (77), "the,principal v a r i a t i o n  contributes  conclusion  fluctuates  base  cells  initials  by L a r s o n  diameter  was  diameter  for  stated  of tracheids  increase  wood  which  tracheid  to  quality.  are primarily  from  of cambial  reference  the importance  post-division  tangential  variation  structural A  little  special  section (3).  As in  diameters  initials  as t h e w i d t h s  cross  reviewed  i n r e l a t i o n t o wood  Tangential widths  (48),with  t h e stem decreases  17.  Investigations indicate wall  that  less  are independent of those  although  these  oping  determined  tracheid, wall  amount o f s u c r o s e , (80).  Tracheid  section  t o a maximum  120).  with  Within  height  latewood  tracheids  thickness (57)  rapidly  from  i n mid-stem  She f o u n d  the second  very  i n r e d pine  increased  thickness  the cells  with  with  decreases  exhibit a  r e f e r mostly tracheid  slight to  wall  (77).  of  Hiller  latewood  of l o b l o l l y  thickness  pine  increased  internode  from the  Larson  (78)  A i t . ) earlywood  wall  age, whereas  i n latewood  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 .  o l d trees  parison  gradually  tracheid  (65,  t o the tree base. resinosa  by t h e  conditions  sheaths  that wall  (Pinus  devel-  cross  thickness  t o about the eighth  that  increase  wall  a  was  age on a  a n d may  earlywood  e n t i r e annual growth  showed  Finally,  growth  These p a t t e r n s  apex, t h e n more s l o w l y  the  each  with  increment, w a l l  sheath  thickness  reaching  that varies with  and, although  taeda L.).  more o r  t o be d e t e r m i n e d  increases  variation i n radial  i n several  (Pinus  appears  thickness  t h e crown.  secondary  tracheid  reaching  also v a r i e s , the variations are minimal  studied  cells  thickness  t o some p o i n t  within  regulating  by t h e amount o f a u x i n  a growth  increase  controlling  Wodzicki(164)  Whereas t r a c h e i d d i a m e t e r  o r photosynthate,  wall  (112)and  two c h a r a c t e r i s t i c s do v a r y  simultaneously - i n nature.  primarily  77,  (77), R i c h a r d s o n  the p h y s i o l o g i c a l processes  thickness  diameter,  of Larson  that  comprise  the outer  (320 t o 400 y e a r s )  wood  formed during  are often  earlier  growth  increment of  thin walled  stages  i n com- .  of growth (52).  18. Within fibril is  o r i e n t a t i o n , and  r e l a t e d to the  microfibril  (109, the  110).  fibre  first  He  was  dimension  that  was  i n longer  shown t o a p p l y  increments  as w e l l  increments, aging  also  so  as  143,  the  (145)  and  of  later  the  T h i s was  f o r the  and is  fibril  angle  decrease i n f i b r i l  that m i c r o f i b r i l length further  are  noted  that  between m i c r o f i b r i l  ones.  The  relationship  from s u c c e s s i v e  growth  f r o m a number o f reflect  a  subsequently  process  (110)  and  cotton  as  fibres  (57)  Likewise, cell  wall  in wall Necesany  thickness,  and  of  similarly thickness there  showed cell  of mutual d i r e c t p r o p o r t i o n a l i t y . an  the  (145).  thickness, (97)  apply  in  fibres  c o r r e l a t i o n between r a d i a l w a l l increase  (see  shown t o  146),  angle.  growth  t r a c h e i d s were d e r i v e d  (145,  thickness,  features  Preston  (micro-  c o m p r e s s i o n wood  i n that, with  of  of e a r l y growth i n c r e -  i n t e r e s t to note t h a t H i l l e r  degree of  by  as w e l l  phloem f i b r e s  found a high  1934  (89),  angiosperms  i s of  as  the morphology  layers  of  It  and  r e l a t i o n s h i p d i d not  abnormal t r a c h e i d s (145),  composition,  l a y e r i n gymnosperms  fibres  tracheids  145).  fibre  micro-  A r e l a t i o n s h i p between  ago  t o random samples  t h a t the  144,  long  the  c e l l u l o s e molecules  of meristem from which  142,  wall  f o r the  short  fibres  to the  as  the  i n the  (145).  cell  reported  observed  greater  ments t h a n  for  to l e s s e r degree the  angle of o r i e n t a t i o n of  fibrils)  i s some e v i d e n c e t h a t  o r i e n t a t i o n i n the  wood f i b r e s was  of  a tree there  He  inverse proportional relationship exists  thickness  and  lignin  content  i n the  fibre.  There  are indications that  related.  Thus  (6) , G r a f f (56), is  and Wheeler  diameter  length  with  have  may  also  be r e l a t e d  review  relationships  exist  and w a l l  between  Simmonds  that  increase wall,  there  length  a recent  wall  these  shows  and  study  by  i n fibre  f o r coniferous  thickness  features  that  dimensions,  hence with  and Recovery  and  and  are i n -  angle.  above  cell  organization;  Composition  both  microfibril  presented  r  Bannan  a n d - H i l l e r , (57), i f  out that  since  them  fibre  of fibre  ( 1 0 9 , 110)  point  are also  demonstrated  In addition,  thickness  c o r r e l a t e d with  Chemical  (54) , H e i n i g  out a r e c t i l i n e a r  of Preston  together,  The  length,  (158),  species.  increasing  length  versely  et a l .  each  Results  considered cell  (50) , H a r l o w  (140) b r o u g h t  species..  and w i d t h  p o s i t i v e r e l a t i o n s h i p between  within  Vorreiter  length  a n u m b e r o f i n v e s t i g a t o r s , among  and M i l l e r  a direct  cell  of  definite  inter-  i n particular  tracheid  composition.  Carbohydrate  Fraction  Wood cellulose, the  major  fibrous lose  of three  hemicelluloses, constituent,  material  i n wood  usually  consists  obtained  lose/,of which  and l i g n i n  constituents, (135).  angiosperms  lignin-free  a l l of the hemicelluloses  The t o t a l  f r o m wood  namely  Holocellulose i s  and i s , by d e f i n i t i o n , t h e  comprising  (128).  main  cell  wall  substance  contain  from  polysaccharides  i n the form 70  and  cellu-  are  of holocellu-  t o 80 p e r c e n t ,  and  20. conifers chief  60  to  75  per  value  of  holocellulose  beginning material  for  by  alpha-cellulose,  in  angiosperms  this  and  40  tion  of  glucan  45  per  and  mannans f r o m  carbohydrate  sample,  are  determination  of  true  20°C  (128,  about  Zobel  The  g  c h l o r i t e , Watson  Leopold  (83)  hydride  reduction.  employed p e r a c e t i c  , 91)  hydrolyzed  A  and to  In  fifth  Larson  162,  well  that  in a  solu-  a  or  homogeneous carbon  as  significant  holocellulose 2 g  of  or  wood m e a l four  used acid  and  Erickson  by  i n which  is calculated  (43)  from  and  sodium  boro-  been used  secondary  xylem  from which some b a s i c  for  fraction  chlorine-sulphate, followed  per  micro-  s i m i l a r carbohydrate (171)  is  163).  addition,  t h e i r monosaccharides,  composition  be  of  term, however,  s e m i - m i c r o - m e t h o d has (78)  to  samples, have been advanced  McElwee (149)  as  (20);  fraction  amounts of  determining  129).  0.5  and  applied  saccharide  for  in conifers  exclusively  (126,  holocellulose  is insoluble  small  a  approximated  cent of  cellulosic  (131).  imply  conifers  as  i s considered  pulp which  holocellulose  161).  (90  that  fractions, requiring  available  methods, r e q u i r i n g  are  the  per  furfural-yielding materials,  similar  Ref.  50  The  i t offers  Glucan,  cent of  does not  S t a n d a r d methods  Meier  to  f o r , i n v a r i a b l y , i t contains  amounts o f  (see  45  as  or  i s that  research.  per  c e n t NaOH a t  a r b i t r a r y , and  dioxide-  e x t r a c t i v e - f r e e wood.  Alpha-cellulose  holocellulose  17.5  strictly  further  to  on  preparation  regarded  wood c a r b o h y d r a t e s . of  based  constitutes  i s customarily  fraction  cent,  the  by  fibres poly-  assumptions  21.  and  chromatographic  literature (123).  on these  Squire  also  determination,  be  A major  applied  aspects  For  yield  example,  with  the l i g n i n .  rigorous  Thus  in  o f n i t r a t e d wood  o f wood  lignin,  present  that  may  the analyst  i f no c o n s t i t u e n t s  use pre-extracted  (20). A  not  and m o i s t u r e - f r e e  arise  from  d i f f e r e n t woods ( 2 0 ) .  that have  variability  n o t con-  extracted  As s u c h , t h e  been that  should  overlooked includes  accounts prefer,  summative  basis,  by  to account f o r ideally  workers  and t o r e p o r t  (137).  i s represented  aims  summation  Many  varia-  on h o l o c e l l u l o s e f o r  and ash i d e a l l y  o f wood.  wood  i s  are inconsistent.  by a summation  holocellulose,  extractive-free  f r a c t i o n a s some data  o f wood  (128) d o e s  of a n a l y t i c a l data  i n which  constituents  sample  cellulose  the published  evaluation  analyses,  i n t h e same  and Bevan  i s no o v e r l a p p i n g  errors  yield  show  there  the  intra-incremental  will  100 p e r c e n t ,  to  for cellulose  i n a n a l y t i c a l procedures  total  chemical-  Squire  it-could  hemicellulose  constituents  tives,  micro-method  by  that  composition  p a r t i c u l a r species  all  i n a thesis  o f h i s t e c h n i q u e was  the Cross  the entire  summative  review of  t o a l l woods.  of chemical  most  a new  as t h e c o r r e c t e d  limitation  tain  a  comprehensive  i s contained  developed  Differences tion  A  and q u a n t i t a t i v e l y e s t i m a t e d  alpha-cellulose meal.  evidence.  extrac-  f o ra l l however,  analyses  because  i n extractive  and  of  on  possible  content  In loses, are  and l i g n i n ,  approximately  wood): and  substances  fers,  analyses  between  many  exceptions  from  that  wood  contains  galactose  listed  o f these  been  chemical  residues  species. than  o f r e a c t i o n wood  varia-  however,  differs  b u t more  cellulose  n o r m a l wood.  46 p e r c e n t  residue  compared i s much  Cross  increased  change  i n lignin  occurs  has a  reduced.  The  based  that  on  loblolly  i n compression  t o 35 p e r cent,  i n total  wood.  analysis, varied  to 35 per cent 28  and more  t o normal  a polysaccharide  and Bevan  from  Tension  wood  a n d Bray- (107) r e p o r t e d  i n n o r m a l wood  no s i g n i f i c a n t  Compression  content  i s instead  Pillow  from  glucomannan  genera;  and x y l a n  a l p h a - c e l l u l o s e , from  and/or  chemical  (136) .  hemicellulose  ently , the increase  elemental  respects  pine  with  ( 1 3 5 , 136)  i n several  amount o f mannose  L i g n i n content  between  coni-  reported.  lignin  residues.  that  composition  than  (20), T i m e l l  Generally,  of  Inter-  (20) a l s o h a s shown wood  i s less  have  less  by B r o w n i n g  relative  tively,  The p r o p o r t i o n  f o r several  and a low c e l l u l o s e  wood.  135). v  cent,  components,  lignin  galactose  25 t o 35 p e r  i n a l l chemical  o f n o r m a l wood  predominant  lignin  (135).  species  The  of extractive-free, dry  p e r - c e n t (20',  Browning  f o r some  preparation,  1 per cent)  (123).  tion  (per cent  methods o f  hemicellu-  (about  i s conveniently  Squire  high  20 f o . 2 5 i s small  variation  of cellulose,  on s t a n d a r d i z e d  40 t o 45 p e r c e n t ,  hemicelluloses  species  The  based  as f o l l o w s  cellulose  pectic  and  conifers, the proportions  pentosans.  a t t h e expense  f o r c o n i f e r s , and t h e r e v e r s e  of  respecApparcellulose  seems t o  operate some  i n t h e . r e a c t i o n wood  evidence  that  compression  wood, q u a l i t a t i v e l y . cellulose  DP  chemistry  Thus,  (degree  wood  of polymerization)  An e x t e n s i v e  chemical  p r o p e r t i e s of compression  review  work  from  normal  (33) i n d i c a t e s l o w  f o r compression  of literature wood  There i s  wood  on a n a t o m i c a l  i s given  by  variations which  aspect  related  i n carbohydrate  are s i g n i f i c a n t (171) a n d B y r d  weight,  loblolly  pine  et a l . outer  more h o l o c e l l u l o s e ,  cellulose general, initial  than  that  Westing  increase  to  the pith,  It  was  wood  and somewhat l e s s  shown by Kennedy  from  content change  and Jaworsky  Zobel  and  that, based  on d r y  one t o t h r e e p e r per cent  wood,  more  alpha-  trees.  In  there  i s a  the increments  beyond  i n increments  1 t o 5 t o 61.6 p e r c e n t  i n increments  25.  Only  further increase  o f 1.7  per cent  tree.  Likewise,  (141)  for  observed  an i n c r e a s e  t o 65 p e r c e n t t h e same  yields  of the 80-year-old  within  species.  i n Cross  the f i r s t  core.  (69), f o r Douglas-fir,  cent  the remainder  rapid close  the juvenile  Cross  a modest  cellulose  diameter,  the.same  from  includes  that  58  and Bevan  t o seven  f o r coniferous  i n cellulose  wood.  contained  wood  topic  a tree  (22)^ r e p o r t e d  and f o u r  that,  across  of juvenile  of juvenile  i t appears  to the thesis  content  because  McElwee  in  and  152).  Another  cent  also differs  Dinwoodie's  fibres.  (151,  o f hardwoods.  increased  and Bevan  from  57.4 p e r  was  from  reported  Wardrop  cellulose  20 i n c r e m e n t s  16 t o  from  pith,  Chlorite also  follow  the above  to  increase  to  the conclusion  fraction  less  gross  the  pith  This  rapidly  than  i n increments  close  to the p i t h  partly  closer  and t h a t  was  and by Wardrop  with  radiata  pine  also  reported  wood  ( 3 2 2 t o 400 y e a r s )  in  were  first  observed per  prevails  They  at that  may  of the older  be.  cell  wall  radiata  for lignin.  across  t h e stem.  species  Hale  percentage  and  with  and  Clermont  i n overmature  contained  this  from  (52) w o r k i n g  t h e same  (D. D o n . ) .  attributed  that  could  thin-walled  to the extremely  contribute  slow  to the differences  variation within  differences  i n growth  i n 1926 by R i t t e r a n d F l e c k  c e l l u l o s e forms  of three  increments  length  with  generally  amount i n v o l v e  reported  i n behaviour  age.  factors  of the total  earlywood,  This  and Clermont  (141),  that  Quantitative  that  cent)  (Pinus  cells.  carbohydrate  increment.  of tracheid  a low a l p h a - c e l l u l o s e  Other  (70).  i n successive  shown by H a l e  Douglas-fir,  o f wood  hemicellulose  of i n t e r e s t i s the s i m i l a r i t y  c e l l u l o s e content  outward,  prosenchyma  (52, 6 9 ) , l e a d i n g  a greater  to the autohydrolysis  appear  to the p i t h (70).  point  wood  be  patterns  percentages  alpha-cellulose may  similarity  growth  Holocellulose  there  A of  trend.  and a l p h a - c e l l u l o s e  that  attributed material  holocellulose  wood  larger  substance  coniferous These  a  t h e growth zone  chemistry  (114).  percentage i n latewood  (up t o  3.4  than i n  woods, and an o p p o s i t e  r e s u l t s were confirmed  They  situation  f o r conifers  by  subsequent  fractions however, than  workers  (52, showed  reported  An earlywood level  123,  of  season.  124,  150,  greater  by  anomaly  earlywood  Squire  minimum  peak  for  while  first-  (72.5  to  and  estimated  between  shown  for  latewood  is  per  site  rate,  and  diameter  gravity (165)  index  of  that  the site  at  at  per  the  cent  wood.  the  seem  index,  in  the  of  yields  three  tree total  i t was is  the  a  at  at -the  .Wilson  to  78  per  latewood  adjacent  (156) , cent,  initiation, had  lower  increments  differences>  than  simi-  occurred  analyses  were  year),  such  increments  latewood  higher  in  of  preceding  present  retain  (77  of  chemical  the  the  to  point  average  height,  which  of  Further  cells  associated  Thus,  from  the  by-Wellwood. and  and  content  breast  workers,  first-formed  at  tissues  quantitative  earlywood  Carbohydrate by  the  holocellulose  cent),  Douglas-fir, six  later  cellulose yield  reported  occurs  Besides  differences  of  not  last-formed  73.5  the  magnitude  in/that  (i.e.,  does  chlorite  lignin)  Douglas-fir.  that,  the  1  corrected  of  in  last-formed  (123),  intra-incremental. lewel;  the  Some o f  r a d i a l - c e l l u l a r • depth .  that  alpha-cellulose  r e t a i n some s i m i l a r i t y  Later-formed  considerable  values  here,  to  Thus,  indicate  161).  organization  to  and  Fleck.  occurs  to  holo-  difference  R i t t e r and  appears  according larity.  f o r both  qualitative  noted.  It  c e l l u l o s e chain that  wood  of  properties height,  shown by  be  and  (161).  influenced  such  Worster  composite.of  length  earlywood  could  was  as  growth  specific and  several  Sugiyama basic  factors,  has a pronounced  conifers  and a n g i o s p e r m s .  that  the width  saccharide ted. that the  much  of annual  amount  does  contents  of cell  wall  wall  layers  effects  this  could  substance  he  notes  concentration  volume  i n  There-  and latewood  to specific  per unit  found  poly-  to density.  i n earlywood  be r e l a t e d  (87)  are elimina-  concept;  i s related  differences  content of  not influence  gravity  confirms  McMillan  of the v a r i a t i o n i n polysaccharide  the observed  bohydrate  specific  on c a r b o h y d r a t e  to this,  increment  (90) a n a l y s i s  different cell  fore,  Contrary  c o n t e n t when  Meier's  influence  gravity  caror  i n t h e wood  (124) .  Estimates in  of Carbohydrate  Relation  to Cellular  Interest cells wall  i n woody physical  stituents  any  studies of  organization,  layers.  fraction)  t o two p e r c e n t )  compositional  data been  average  has been  Thus,  Thornber  weight  from  wood.  related  (amount o f  very  little.  (weight  to  Some basis)  and r a y parenchyma  f r a c t i o n a t i o n of the  In only  one i n s t a n c e  to the weight  of single  con-  and i t s r e l a t i o n  the percentages  and N o r t h c o t e  and l e n g t h  factor  studied  by q u a n t i t a t i v e  derived  of individual  on d e t e r m i n i n g t h e  The w e i g h t  (98 t o 99 p e r c e n t )  holocellulose  unit.  mostly  of individual cells  element  tracheids  examination  and d i s t r i b u t i o n o f c h e m i c a l  ( 1 0 , 1 0 4 , 105) r e p o r t e d  vertical  (one  Elements  t i s s u e has centred  particular cell  and Weight  i n the intensive  i n the wall  carbohydrate  Fraction  of a  (13 4)  have  cells  i n four  total  have  biological  estimated the species  consisting that  of three  angiosperms  t h e mean w e i g h t  and one c o n i f e r .  of the c e l l  increased  as  3 50 t o 930 p e r c e n t  i n the four  It  was  at a l l stages,  fer  concluded  that,  were.considerably  No  attempt  in  cells  was made  to relate  fibres.  Weights  a Mettler  length  with  tracheids filament  was  cell  were based UM7  examined  those  from  the observed  by  from  them.  the coni-  the angiosperms.  weight d i f f e r e n c e s  first  balance.  An  density  and expressed  ingenious  attempted Weights  the species  wall  w e r e made  on measurement  balance,  (coarseness).  Neither  than  a g e b y a s much  the cells  of tracheid weights  i n connection  using  species  found  to length..  Estimates (166)  heavier  with  They  method  by Skene  reported  employed,  by  Yiannos  of commercial  pulp  o f a group  fibres  of  as w e i g h t  per unit  of weighing  single  (121) u s i n g  ranged  from  a  quartz  0.04  t o 0.1 y g .  nor the tracheid composition  are  available.  Weighing property sions. also  and r e l a t i n g  has s e v e r a l advantages I t i s not only  the effects  the" a m o u n t property, ness'  cells  of cell  of cellulosic i.e., fibre  and i s d e f i n e d  expressed  much  easier  wall  measuring  composing  p e r 100 m  and c a l l e d  In view  dimen-  but includes o f t h e lumen,and  the c e l l s .  i s usually described per unit  morphological  lateral  and q u i c k e r ,  as t h e w e i g h t  t o 'dg' ( 1 3 2 ) .  to a  thickness, the size  material  weight,  i n milligrams  abbreviated  over  results  as  length; a  This 'coarse-  i t i s  decigrex,  of i t s important  effect  on  28.  many p r o p e r t i e s (24)  and  been  a method  described  A in  wood,  section 17).  and  nique  ness  with  is  tree has  of  upon  but  formed  that  per Such  only  specific  wood  pentosan  content  known and  angiosperms teristic to  of  as  the  length  a  length,  of  weights,  differ ray  some  his  co-workers  prosenchyma,  time  conifers.  chemical is  feature their  (1,  cells  for  and  Britt  this  fibre  fibres  be the  tech-  was  per  pulp  weigh  unit  more.  centre  on  a  coarse-  weighs  studies  (16,  coni-  there  from  cross  of  a  samples  more  than  justified  on  the  basis  tracheid  morphological  earlier.  the  cells  in  older  on  can  of  western  that  increases  longer  conclusion  by  mm  indicating that,  grows  based  coasreness  following  increase  have  (132).  square  various  studies  tree,  tree  su-67  fibre  per  w e r e made  such  the  fibre  unit  discussed  Not  Perila  from  of  samples,  described  on  inter-relationships within  properties  been  that  ages,  irregular  age  T234  fibres  measurements  'coarseness'  for pulp  g r a v i t y , was  shown  fibre  Method  of  thus •Clark\(24),  ones.  known  number  somewhat  known  of  determination  various  I t was  outward; shown  for  coarseness  fibre  also  Suggested  specific  at  the.concept  i t s measurement  the  increasing  a  shorter  for  method  the  (16).  length, It  new  species,  progressive  paper,  i n TAPPI  based  Fibre  ferous  of  (1,  but 55,  103,  55,  Their the  133, 104, data  to  142).  the  136). 105)  Recent  indicate  high  The  higher has  studies  this  that  cells,  xylan  of  e n t i r e wood  confirm  parenchyma  unusually  composition  133,  compared  (103,  of  chemical  for  the as  content.  of both  charac-  compared  Effects and  of  Fertilization  Chemical  of  spruce  of  fertilized  Posey is  (71) , i n  and  pine  presented  stands  results  slight  change  in  size ies  and  shape  reported  the  of  so  (117).  in  cells  far  are  of  reduction  cent  than  compensated  71,  117).  Any produced that  the  would  have  brought  decrease  yield  perhaps  f o r by  is  to per  less on  about  the  be  in specific  both  properties by  but  of  resulting  (79).  by  Lee  (81),  information  of  by  less  and  that  the the  product.  changes  that  Stud-  the a  maximum  production  the  and  a  raw  paper  in either  (44,  i f  industry  Additionally,  change This  more  material  importance  reduced.  effect  in  i t is usually  greatest  be  is  to  a  consistent.  gravity  pulp  and  Changes  i n volume  the  fertilizers  response  g r a v i t y of  volume w i l l is  such  fertilizing  increase  utilized  wood q u a l i t y  Larson  ratio.  specific  be  characteristics  provided  growth  and  occur,  will  by  main  thesis.  minimal  appreciated,  the  been  increased  can the  unit  wood  a p p l i c a t i o n of  s e n s i t i v e to  fertilization  material  since and  by  this  indicate that  most  per  an  also  Consequently,  in  the  i n assessing  earlywood-latewood  characteristic 10  have  instances,  forest  on  t r e e s were d i s c u s s e d  in detail  most  summarized  stands  Problems  surveys  Sastry  paper,  forest  species.  literature  In  of  coniferous  (108)and  not  Properties  a recent  fertilization  of  Detailed  Wood  Composition  Klem effects  on  in  could the  density be  proportion  30. of  latewood  cell  f i b r e s i n t h e raw m a t e r i a l ,  morphology,  and p o s s i b l y  type of product being advantageous the less be  produced  of deleterious.  quality of this importance,  material  not because  altered but, with  would  n o t be l i k e l y  cell  current  or through  weight.  Depending  t h e changes  could  The i n f l u e n c e when u t i l i z e d the strength grading  i s of  of the product  p r a c t i c e s , these  t o a f f e c t lumber grade.  a l l fertilizer  i n obtaining  (92, 93).  the basic  experiments,  Consequently,  that  stands.  The  same  authors,  differential  as w e l l  properties  cal  a  and growth  rate  (117)  effects of f e r t i l i z e r  been  growth  to learn  more  and q u a l i t y o f  fertilizer  treatment  gravity of loblolly  as Posey  o f wood.  of f e r t i l i z e r  t o r y work by S a s t r y tial  i n wood  F o r e x a m p l e , .^Zobel et. a l .  (108),  effects of f e r t i l i z e r  properties  formulations  machine  have  however, have  i ti s imperative  c e r t a i n l e v e l s o f NPK  d r a s t i c e f f e c t s on s p e c i f i c  initial  possibly  factors a f f e c t i n g the quantity  had  of  changes  a s u b s t a n t i a l and c o n s i s t e n t  wood grown i n f e r t i l i z e d found  factors  e f f e c t on lumber g r a d e ( 6 6 ) .  Not  (173)  could  on  could  I f and when  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  about  be  as lumber  quality  response  either  changes  o f s t r u c t u r a l lumber becomes i m p o r t a n t ,  successful  upon t h e  of such  grading  significant  changes i n  Posey  the  wood.  existence  treatment depending  also  found  brought greater compared  showed  pine  that  certain  difference  to others.  i n wood  Earlier  indicated the p o s s i b i l i t y  on  explora-  of differen-  c o m p o s i t i o n on p h y s i c a l and mechani-  properties' of Douglas-fir  wood.  The influence  on  possibility, older  tors  relating  real  e f f e c t s of  reason  why  to  fertilizer  be  nutrient  western  larch  results  indicate  such  cell  as  Murphey  may  have  Such  a  et  al.  differences  for  this  is part  failed  to  growing  (Nutt.))  selected  fertilizer  the  elicit  an  influence  could  employing  experiments seasons,  in  using  seedlings.  of  the  of  anatomical  number  fac-  confound  conducted  two  dimensions,  may  of  several  experiments  (96)  in  trials,  confounding  laboratory  cultures,  and  field  studies  micro-site  (Larix occidentalis  size  in  application;  response.  Thus  and  trials  controlled in  seedlings. controlled  the  that,  i.e. , in  environment  growth  possibly  trees,  some o f  increased  exists  Their features  tracheids  per  2 10,000 per  ym  , resin  resin  canal,  canal as  Changes tion  have  ficant  been  Lambert  (44),  cellulose zation. showed (173). zation more  in  but  content  a  has  than  earlywood  chemical to  was  a  found  i n young trend,  experimental  been that cells  shown of  to  with  their  number and  very in  l i m i t e d degree.  lower the  wood  the  the  that  higher  not  lignin  and  fertiliyield  significant  expected.  greater content  signi-  alpha-  after  production  the  and  cellulose  c h a n g e was be  fertiliza-  Erickson  formed  pine,  r e s u l t s would  so  by  cells  properties.  Little  holocellulose  loblolly but  epithelial  following  Douglas-fir  in  of  mechanical  composition  increase  latewood,  and  physical  slightly  decreasing These  as  reported  Likewise, a  well  studied  c h a n g e was  diameter  of  Fertili-  earlywood  number would  of decrease  cellulose clusions on  percentage  are i n general  increase  abies  lamella  growth  particularly properties due  in  et a l .  paper  burst  rate  were  produced  tics  due  collapse form  such  as f o u n d  fibre  as r e v e a l e d fibre  Thus,  Furthermore,  i n high  tensile  i s a dominant  r e s i s t a n c e of paper,  long  obtained  characteris^„of  proportion,  resulting  present  highly  i n strong  strengths  producing  to  i n for-  factor controlling  fibres  large  process,  they  paper. .  explained  uniform  and b u r s t i n g  higher  bulky  fibres  i n high  low i n o p a c i t y ,  areas,  Wood  c a n be  i n the conversion  contact  was  i n wood  wood  rate  reported  and l e s s  thin-walled  i n fertilized  of surface.  length  changes  paper  than the  trees.  characteristics  inc-  growth  fertilization  a denser  of  that  (51) h a v e  i n tear  trees.  on wood q u a l i t y ,  reported  unfertilized  sheet,  to fibre  the effect  i n Sweden  o f t h e known  (Picea  i n fertilized  the increased  and lower  con-  (42) f i n d i n g s  spruce  i n Finland  I t was  produced  i n paper  and smooth  Similarly, tearing  from  also  dense  studied  after  to f l a t t e n e d ribbons  effective bonds,  wood  These  on t h e i n c r e a s e i n  cells  and spruce  Studies  material  a compact,  mation,  of latewood  to fertilization.  diameter,  (88) d a t a  strength  on t h e b a s i s  of f e r t i l i z e d  a f f e c t e d by  from  and t e n s i l e  Changes  content  and p a p e r .  little  fertilization  readily  Erickson's  of pine  f o r pulp  corresponding after  agreement with  (64) h a v e  to fertilization.  that  of lignin.  a n d Malm's  thickness  Jensen reased  that  i n the l i g n i n  (L.) K a r s t ) ,  middle  and i n c r e a s e  paper  (13). the with  the  highest  area  the  Both  bulk  tearing  shearing  tracheid  and  coniferous the  vigor  to  Extremely  growing  tial  the  the  often  and,  may  1  s h o w no  response  so  f a r are  f o r by  l a t e w o o d ) may  to  changes  could to  cause  an  that  play  anticlinal  a  large  (13).  with  the  to of  increasing  response  be  related  the  to  with  the  growing  fertilization.  to  (up  trees  their  radial in this  to  50  of  rapid  length  division. the  cambial Bannan  means by  but  which  some  related  wood.  i s expected  increase  number o f  per  response.  gravity  fertilized  poten-  thickness  density  in specific  in cell  very  wall  in  density  i n latewood percentage;  some p a r t  d i v i s i o n s are  density  decrease  The  most p a r t  changing  of  low  to  close  indi-  environment.  (51), while found  to  of.the  producing  generally  f o r the  i n the  pattern  are  of  particular  i n t e r p l a y of  i n density  trees  a decrease  increase  change i n t h e i r  level  the  i n chemical properties  i s associated  that  appears  however, were changes  Likewise, this  are  changes  (mostly  study,  stated  which  increase  have i n d i c a t e d t h a t  no  some e x t e n t  initial  trees  studies  In  be  finally,  most vigorous  accounted  to  genetically controlled ability  a moderate r a t e The  over  i n t e a r i n g paper  fertilization  slow growing  changes observed cent)  increase  tree,  respond  at  density.  to  the  'starvation.wood  distributing  (29).  trees of  by  involved  summary, i s can  wood p r o p e r t y , vidual  forces  porosity  length  In  resistance  i n growth, initials (8)  since which due  showed  multiplication  of  fusiform  plicative initials  i n t h e cambium, a  factor  increase  cell  This  ( 7 1 , 108,  the be  initial a large  fertilizer 'individual ment  i s achieved. influence hence,  amount  growth,  concept  length  should  of  of derived  cause  i s supported  as s t a t e d  property  by  Zobel  effect  by  which  of multiderived vascular  would  a drop  cause  i n t h e mean  t h e work  of  several  response'  of the tree.  suggesting may  e t al_. ( 1 7 3 ) , t h e r e i s  of f e r t i l i z e r  of individual tree  treatments,  of improved  the length  as f e r t i l i z a t i o n ,  of a differential wood  Frequency  117, 141, 1 7 3 ) .  Finally, indication  such  i n radial  length.  authors  cells  d i v i s i o n s would  elements; an  cambial  become  strains of  Also,  very  trees.  there  difference  genetic  depending appears  to  i n response  to  interaction.  important  on  This  i n the develop-  CHAPTER I I I  MATERIALS  Selection  of  tracheid  selection ment  of  a broad  range to  o f wood  length  and  more  The  radial  of  and  (Picea  species  of  coniferous  a  sempervirens white  pine  that  the  material.  might  be a  pattern  was  based  was  this  oVer  i n tracheid  sampled  i n one  available  in this  disk  measure-  so a  that  reasonable In  length disk,  order from  at  (Pseudotsuga from  has  another  been  the  objective,  made,  interpreted  range  for  on  general statement.  were  a  criterion  Within  tissue  variation  which  as w i d e  length  500-year-old Douglas-fir  menstudy.  described  by  (70).  sugar  pine  sitchensis  three species (Pinus (Bong.)  representative  (D.  so  include  tracheid  approach  Pinaceae,  families  to  o f wood  points  Franco)  Warren  sampled:  spruce  a  density  From were  t h e work  several  (Mirb.)  Kennedy  was  the w i t h i n - t r e e  height,  ziesii  material  closely  was  possible,  from macerated  from  to bark,  stump  objective  taxonomical sorting  include  pith  primary  l e n g t h s as  conclusions  METHODS  Material  The of  AND  were  Don)  (Podocarpus  Carr.)  genuss  from  separate  genera  lambertiana Dougl.),  included;  Endl.)  from  from  and  Douglas-fir.  each  of  t h e s e were Taxodiaceae,  d a c r y d i o d e s A.  Rich)  Sitka  the  s i x other  redwood New  from  One  (Sequoia  Zealand Podocarpaceae,  eastern  redcedar  Pacific  yew  cow's-tail taxaceae,  (Juniperus virginiana  (Taxus pine  brevifolia  Araucariaceae.  pine  wood w e r e  Douglas-fir  (119).  Two  were  consisted  which  generated, (NH^^SO^)  zation  fertilized  one  growth  formed  nine  28 g r o w t h redwood, was  available  University  with  and a l l seven  pine, from  gravity  were  and  New  of a disk,  was  Experimental Research  A  and f i v e  Douglasincluded  increments  applications.  nine  genera, A  Wood m a t e r i a l pine,  wood  of  and  total  of  of sugar  and e a s t e r n  pine,  redcedar  Forestry,  sample  o b t a i n e d from Forest,  decrease  fertili-  of a  families.  small  treat-  application.  sampling  included  Zealand white  Columbia.  and  following  the xylarium of the Faculty  of B r i t i s h  The  ( N H ^ ) ^  (Urea)  tree,  coniferous  sampled.  fertilizer  b e f o r e and a f t e r  t h e two  plan  study  study m a t e r i a l .  In this  sampling  from f e r - .  of three  an i n c r e a s e  and a f t e r  Douglas-fir.  another  NPK,  from  pronounced  of  cores  from  each  Cephalo-  Sweet)  a tree  before f e r t i l i z a t i o n  entire  taxus , i n t h e form University,  this  variation  NH^NO^.  increments hoop  one t r e e  from  from  i n s i x consecutive increments  with  coincident  species,  available  specific  weight  increment  The  from  Taiwan  showing  of increment  respectively, i n wood  Taxaceae, Hay.)  of formulations o f Urea,  traced  fir  was  Cupressaceae,  cunninghamii  included  from  selected  tracheid  was  trees  increments,  ments  Finally,  also  from  two i n c r e m e n t s  i n t h e form  tilized  (NPK,  (Araucaria  In addition,  Material  treatments,  from  (Cephalotaxus wilsoniana  and hoop  compression  Nutt.)  L.)  of  Cephalo-  National  i n Taiwan.  The  Taiwan . -  remaining the  material  author,  Forest,  near  Vancouver  given  Wood  Haney  i n Appendix  excised  carefully  the  increments  yield  then  into  latewood  South  Central  wood  characsamples  growth  or increment  into  and  smaller  packets;  each  the species,  prepared  increments cores,  latewood pieces,  i n stainless steel  small  also  wood  wire  packet  increment  Separate  packets  to determine  the  then  portions; with  screen was  labelled  number,  and  o f some o f percentage  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 .  were  cellulose pieces  Research  o f t h e s e wood  earlywood  fractions.  Holocellulose pieces  disks  dissected  separately  were  by  Preparation  Selected  blocks,  tag to identify  and  Valley,  length  Skeleton  i n t o whole  a n d made  a metal  earlywood  was  bundled  (150-mesh) with  wood  separated  Columbia  d e s c r i p t i o n of gross  procedure:  from  of these  portions  River  Columbia  1.  and T r a c h e i d  General  each  and Robertson Origin,  British  of British  and measured t r a c h e i d  Sample  were  collected within  the University  Island.  teristics, are  from  was  extracted  tracheid  with  reduction,  (83).  method  were  following  i s reported  fibres  with  93  Packets  containing  (V/V) e t h a n o l - b e n z e n e ;  i n peracetic  borohydride  holocellulose  2:1  skeletons  ( i n packets)  This  preparation:  obtained acid/  by  with  t h e method  to give  t o 100  holo-  t r e a t i n g wood subsequent  described  virtually  per cent  wood  by  intact  recovery  sodium Leopold superior  of major  38.  polysaccharides included  1.  three  The  a n d minimum  treatment  by  temperature  water  packets  the  flask was  glass  of f i v e  cycles  The  holocellulose tubes  and  2.5  total  temperature  i n h o t water  cycle  (30  (3 o r 5;  in distilled acetone.  then  70°C),  minutes,  see below) water  After  placed  evacuated,  drawn  presence  earlywood  test  o f wood;  o f wood;  and  and t h e  air-drying,  i n a suction a new  the  batch  flask, of r e -  into i t . borohydride  9.5), the d e l i g n i f i e d ,  defiberized  by  rod against the wall  for  in  were  was  were  acid  acid  70°C).  (3 h r , pH  pieces  of 30  of a peracetic  A t the c o n c l u s i o n o f the sodium tion  total  soaking  removed w i t h  agent  A  p e r gram  were washed  the  the  procedure  a number  5 g of peracetic  A t t h e end o f each packets  into  a c e t a t e p e r gram  32 m l  followed  divided  consisting  (i.e.,  o f sodium  volume  3.  was  cycles,  treatment  2.  The  steps:  minute  g  carboxyl groups.  of hot  was  tracheid  gently rubbing of a glass  wood  with  beaker  a  i n  water.  needed  delignification  soft,  reduc-  f o r latewood  as d e t e r m i n e d  s k e l e t o n s thus  containing distilled  water. •  by  and  three  physical  o b t a i n e d were  cycles checks. stored  Alpha-cellulose quantity further  preparation:  of the holocellulose reduced  following  Approximately  f r a c t i o n , from  to alpha-cellulose  TAPPI s t a n d a r d method  skeletons  T203 o s - 6 1  accommodate m i c r o - a m o u n t s o f m a t e r i a l . obtained  f o r holo-  separately skeletons  f o r earlywood  i n test  stages  holo-  tubes  containing  allowed  measurements  r e p r e s e n t more c l o s e l y any as  possible  effects  f a r as p o s s i b l e .  which were  intended  (holocellulose) before  alkali  were processed  standard tity  lignin cent,  for lignin:  method  T 2 3 6 m-60  by B e r z i n s  c o n t e n t s were respectively,  samples  Tracheid  i n wet  condi-  tracheid and  i n the tree. were  were those  determination. into  At a l l  condition  to drying  exceptions  kept  done so t h a t  state  due  Lignin  and a l p h a - c e l l u l o s e  of pulp  II.  avoided samples  These  t h i n sheets  Thus,  pulps  and a i r - d r i e d ,  treatment.  Test cellulose  of shrinkage  for yield  to  are given  water.  on g r e e n  their natural  was  yields  i n Appendix  pulps were kept  be b a s e d  The o n l y  (131) s c a l e d  fractions  T h i s was  the  essentially  Converted  distilled  to dry.  could  sample,  a f t e r washing, were a l s o  and a l p h a - c e l l u l o s e  and n e v e r  length  and l a t e w o o d  of alpha-cellulose,  stored  tion  and a l p h a - c e l l u l o s e  each  half  examined  content  s a m p l e s was  i n some o f t h e determined  (130) as m o d i f i e d (12).  equal  The  f o r holo-  by  TAPPI  f o r reduced  quan-  t e s t showed  to or less  than  1.53  that  residual  a n d 0.29  and a l p h a - c e l l u l o s e ,  (see Appendix I I ) .  holo-  i n the  per  40.  Analysis  of  1.  Tracheid  Sampling  The relate were  source.  In  be  obtained  study  such  tracheid is  not  fers,  length  Stein's  to  where  already  i t is  reliable  from  two  of  groups and  been  mean;  10  latewood  establish  the  two  between  such  of  as  a wide  the  f i b r e s each, regions  of  within  (125),  are  effort  (118)  taken  test  of  the  at  tracheid of  sample  of  sample  would  help.  as  to  Previous  that  from  by  time  so  relationship.  increment,  size  the  size  indicated  There-  little  within  random  coni-  workers.  c a l c u l a t i o n of  desired  the  to  in  function  species  several  determine  s i m i l a r study  for  range  primary  i t is  could  However,  within-tree  by  research to  of  c h a r a c t e r i s t i c s , namely  and  procedure  sample  'norms'  increments.  rather  of  determination  included  established  essential  a  to  tracheids  Calculation  defensible  Moreover,  maximize  estimates  experience  r e l a t i o n s h i p the  sampling  those  sample  to  a  a l l  to  irrespective  d i f f e r e n t o r i g i n s , the  procedures  i n order  Initially,  determined.  were  weight.  and  i n v e s t i g a t i o n was  population,  such be  increment  and  this  fraction within  from  two-stage  available,  wood  this  patterns,  fore, standard  get  examine  objective  estimate  have  Still,  one  statistically  tissue as  to  needed a  of  i t s length.  r e l a t i o n s h i p between  tracheid length  for  lengths,  to  overall  to  i f the or  to  belonging  number  number  increments a  as  objective  weight  order  replication sample  criterion  primary  tracheid  treated  Skeletons  at  the  least early-  provide  a  satisfactory standard  number  with  reasonable  the  above  analyzed  from  one  particular  dized  approximately  in  tracheid  up  to  40  2.  length  for  tubes  tracheids a  Care  in was  Tracheid by  was  the  the  has- a v  of  of  a  and  standar-  cases  the  variation  where  n u m b e r was  increased  a  petri  the  These  from  help were  water  d a m a g e was  each  dish; of  even  then and  a  done  the  labelled  randomly  selected  fine  of  tweezers, to  cover  applied.  to  the  Images  screen  their  glass  calibrated +  up  at to  56.X  and  4.9 8 mm  glass fibre  the  probe wheel.  0 . 1 0 4 2 mm  Tracheids  and  using  transferred  recorder.  p r e c i s i o n of  wet of  +  condition, fibres  are  measured  instrument  0 . 0 7 1 2 mm  length  glass  tips.  lengths  This  a  were  at  82  X  measured  magnification.  noted,  from  to  length  a  increments  was  fibre  of  for  tracheids  (increment)  in  aid  separately  of  w e r e made,  opaque  As  this  number  measurements  an  higher  some  material  with  no  on  the  the  distilled  that  magnification. using  mean  length  of  length  projected with  the  sample.  microscope.  taken  use  great,  picked,  drop  the  sample In  transferred  were  a  reasons,  20.  was  portion  binocular  slide  that  Tracheid  A test  of  deviations.  For  at  estimates  a minimum  earlywood each  of  and the  of  10  tracheids  latewood  for  each  wass. m e a s u r e d of  the  h o l o - andv a l p h a - c e l l u l o s e  selected pulps.  42.  Alpha-cellulose length  o f up  tracheid  at  t o 15 p e r c e n t  skeletons.  individual cent  tracheid skeletons,  NaOH  tracheid  This  in relation  was  Length  skeletons  petri  the reduction  treatment  of h o l o c e l l u l o s e fibrous  cause  swelling  shortening factors,  3.  Tracheid  these  hours.  Later  its  fibre,  humidity for  24  were  (CTH)  ratio..  17.5 p e r  minutes treatment  alkaline  could  i n lattice  be  and  simple  expected thereby  mechanical  (160).  the length,  labelled placed  i n vacuum  each  plastic  (-60°C  and  that  Subsequently  (73 + 3 . 5 ° F ,  50 +  dryer  one hour  was  and  Model  f o r four sufficient each  temperature RH)  Several  (Virtis  the boats,  2%  was  boat.  5 u vacuum)  transferred to a constant room  tracheid  weigh  i n a freeze  showed  fibres.  with  and  conditioned  hours.  Tracheids then weighed  f o r which  individually,  ultra-microbalance. quartz  then  experience  dry individual  the  The  skeletons  by  a  weight  were  and d r i e d  length.  explainable  Poisson's  to a separate  boats  by t r e a t i n g  f o r 75  and a f t e r  t o changes  determining  10-100)  to  (21) d u e  i . e . , reverse  transferred of  i n tracheid  of the tracheid,  After  dishes,  measurements b e f o r e  in  to holocellulose  of holocellulose with  confirmed  to  a shortening  further verified  s o l u t i o n i n separate  20°C.  showed  filament  lengths  using  Briefly,  (20 y d i a m e t e r  had been  a specially  the balance and  6.35  cm  determined designed  consists (2 1/2  were  quartz  of a  thin  in.) long)  firmly  fixed  on  free  the  deflection is  at  one  end.  end  and  i t s weight  of  the  filament  s i m i l a r to  the  loading  end.  The  to  yg,  14  0.03  yg,  balance and  1  which  commercial  syringe,to for  is  possible, cover  with  of  then  photometer then  by  measuring  a microscope.  The  c a n t i l e v e r beam  at  has  range  a weighting  times and  higher  than  consists  of  and  a  in position.  to  and  to  protect  was  the  Na^O^)  dissolved and  the using  absorption with  gram  free  by of  in  of a  a  (yg)  air  of  — 6  the  the  filament  with  a  currents.  below.  balance  These  known v o l u m e  optical  =10  of  of  opening  i t from  the  end  plunger,  is its  c o l o r i m e t r i c method,  density  Unicam  maxima the  present  hypodermic  f l u o r e s c e i n were  optical a  0.06  within  i . e . , the  the  i s given  use  achieved flakes  the  from  of  closed  d e f l e c t i o n measured.  standards,  1 micro  the  ml  loading  cover  ultra-microbalance  in  20  The  made  weighings  placed  principle the  that  a mount,  arrangements  step  is  ultra-microbalances.  t o make  compared 1  determined  filament,  c r y s t a l s or  at  weighed  open  c o l l e c t e d and cent  and  be  left  This  filament  appropriate  were  off  crucial  small  (1 p e r  the  to  reproducibility (precision)  case  filament  the  calibration.  the  10  about  during  A  which  is  balance  cut  a  developed a  the  glass  diagram  of  provides  protect  holding  syringe  so  tracheid  with  electrobalances  The  A  The  493  SP  800  and  density  g  loaded  solvent  measured  Recording  460 of  m.U a  on  c r y s t a l s were  of  was  in  against Spectro-  Results series  of  44.  1  2  6  Cross s e c t i o n o f the Ultra-Wicrobalance.  1.  glass plunger  4.  p l a s t i c n e e d l e mount.  7.  radium  solutions tolerance flection it.  range  5.  the strengths limits.  of the balance.  against  (118,  Once  filament)  may  i n d i v i d u a l (+ 0.03  121).  was  plastic  6.  lock t i p  cover g l a s s l i d  design Weights  was  was  weight  a graph  close  showing  of material  sample  weight  linear  over  used  be weighed  ug).  de-  placed  and v e r t i c a l  the entire  t o weigh  on  dis-  useful  tracheids.  at the rate  reproducible Further  and c a l i b r a t i o n o f some  known w i t h i n  c a l i b r a t e d , t h e balance .(using  t h e r e s u l t s were  construction,  were  The f i n a l . r e s u l t  of the filament  same q u a r t z  same  quartz filament  o f which  of the filament  minute;  3.  foil  Tracheids per  open-ended g l a s s tube  The r e l a t i o n s h i p between  placement  the  2.  when  details  o f o n e o r two reweighing  of the balance  are discussed  of the tracheids  the  elsewhere  (1 t o 2 u g ) a r e  comparable UM7  (+ 0.15  balance,  yg)  as d e t e r m i n e d  Correction tracheids room.  thus  follows:  water  were in  r e m o v e d by  numbered  t h e CTH  which then  they  per  cent.  are  given  were  ovendried.  condition,  This  thin  was  done  "pulp  stored  of  fibre,  ( i n t h e open) following  weights  to the nearest  of the pulp  number  sheets"  reached,  tracheid  as  and  sheets  These  CTH  samples  were  0.01 examined  i n Appendix I I I .  Analyses  Several proposed  different  contents  of the  the holo-  w e i g h t was  basis,  of  of the required  and k e p t  Individual  to moisture-free  from  into  suction.  a constant  The m o i s t u r e  Statistical  the  made  for identification, until  adjusted  was  a t t h e EMC  weight.  selection  applying  room  a Mettler  The w e i g h t  i n ovendry  the extra material available  t r a c h e i d s f o r weighing)  and  that  the weight  (after  using  content:  to the o r i g i n a l  pulps  by  checks.  represents  to express  were'made  alpha-cellulose of  by  obtained  f o r moisture  obtained  In order  adjustments  to those  analyses  hypothesis  were  conducted  i n order  and i t s a p p l i c a b i l i t y  to  t o woods  test of  origin.  Multiple interrelationships increments association  regression analyses within  (earlywood between  and  were  used  to  determine  and between s p e c i e s , and w i t h i n latewood);  tracheid  length  to assess and weight;  the measure o f finally,  to  develop  regression  least-squares zero  of  models,  intercept)  required  equations  were  a stepwise  multiple  ferences  conditioned also  within mature  earlywood  and l a t e w o o d ;  trees.  Covariance  dependent sets  (Tracheid  'Y'  between  (Tracheid  of tree  normal  wood  and t o b r i n g  to test  dif-  formed a t  growth;  and  between  compression  o f wood  from  to adjust  Weight)  equation  and Smith ( 7 3 ) .  stages  used  (with  an  b e t w e e n wood  the behaviour  to the  technique  from  by. K o z a k  species;  of the corresponding  Length)  models  were"employed  i s a technique  variable  of values  procedure,  and overmature  and t o c h a r a c t e r i z e  In addition  The r e g r e s s i o n  of -covariance  and between  f i t .  regression  as d e v e l o p e d  juvenile,  wood;  used.  elimination  variables,  Analyses  of best  fertilized  means  of the  for differences i n  independent  variable  out differences,  i f any,  'X' within  2 or  between  compares Its  or  regression  purpose  should  the groups  noted  equations  i s to determine  be u s e d  above.  f o r each  f o rparallelism  whether  population  a l l of the populations  The a n a l y s i s  could  separate under  and  coincidence.  regression  study  be d e s c r i b e d  employed  and/or  by one  equations  i f some  regression-  equation .  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 of 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 at 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 Statist 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 , S e p t e m b e r 7-11, 1970.  CHAPTER  R E S U L T S AND  The of  tracheids  that in  present in  species  matter, with  tracheid  attention  a)  within  focussed  establishing  of  procedures  trees.  a  (holo-  and  in  between  a  basic  biological  aid  of  to  test  the  and  yield  cell  characteristics in  also  provide  heid  length  improvement  means for  on  of  observed  -for  differences and  or  between  carbohydrate that  may  coniferous  tracheid  such  a  exist  of  woods,  concepts, example  length;  basic  and  relationship  fertilization  on  Douglas-fir.  coniferous  re-emphasize  the  species  importance  s e l e c t i o n of i n d i v i d u a l s w i t h i n  work.  nor,  species;  effects  relationship for to,  of  morphological  based  wood  fundamental  test  considerations,for  usefulness to  amount  coniferous  respect  a  origin  i t s purpose  alpha-cellulose)  with  Such  to  description  relationship  relationship for  practical  carbohydrate  Rather,  quantitative  fraction  evolving  with  on:  and.the  and  concerned  d i f f e r e n t taxonomic  length  from  c)  not  tracheid  within b)  of  DISCUSSION  is  development  weights  are  study  IV  species  could of, in  tractree  48.  Relationship  1.  cellulose measured  for  Tracheid  sion  for for  for  Only  wood  its  cellulose  estimate  data  wood a  from  (787  holo-  alpha-cellulose skeletons)  the  in  the  nine  increments  and  from  tracheids  latewood,  values, trees,  i n Tables  I  radially  contained  'normal wood'  fertilized  given and  were of  II.  across  the  i n Tables  (excluding  trees)  was  selected.  are  taken are  species  examined  r e l a t i o n s h i p between weight  addition,  the  the  were  amount  (within  bohydrate  III  compres-  considered fibre  of  fraction,  relationshipslf:or both  examined  separately.  holocellulose is  reasonable  when  skeleton  whereas  limits)  of  An  that  the  holo-  and  alph  advantage  i t provides  total  an  tracheid  alpha-cellulose relates  to  car-  the  basi  component.  Individual tracheid  weight  with  regression  the  weight  length  skeletons  knowing  skeletal  and of  Weight .  length.  In  of  747  Douglas-fir the  and  individual  and  each  and  Length, and  species  1,534  earlywood  two  establishing and  length  weight  IV.  of  skeletons  separately  and  total  values  disks,  Tracheid  Variation within A  Mean  Between  and  species relationshipsobtained length  are  shown  r e l a t i o n s h i p s of  i n Figures  1  best  It  the  c o r r e l a t i o n s determined  using  the  linear  least-squares  r e l a t i o n s h i p , by  significant,  accounting  for  66.4  either  to  94.8  fit. the  to  9,  84.8  along  i s seen  curvilinear  method, and  between  are to  that  or  highly 98.6  per  49.  cent  of v a r i a b i l i t y ,  skeleton The  weights,  variability  cellulose related  than  respectively,  accounted  to the fact  the s k e l e t a l  be  less  f o r by  that  the length  length  i s higher  could  be  alone.  f o r alpha-  Possibly this i s  alpha-cellulose,  There  factor  being  more  basic  framework,  two o t h e r  should  possibilities, ~  being:  The s w e l l i n g and s h r i n k a g e due  to strong  alkali  some o f t h e m i n u t e lationships 2.  The  strong  as  observed  from  length  from  cribed several  pith  This wood  workers  composition influence  removed  isolation  one t h a n  (131),  that of  length  and  1 t o 9 and t h e s t a t i s t i c a l f o r each  i s of importance  bark,  Results  suggested  o f wood  have  and  tracheid  carbohydrate  towards  previously.  occurred  i n length-weight r e -  fibres;  are highly significant,  and g r o s s  that  holo-  (83) .  Figures  examined.  may  of alpha-cellulose  relationships.obetween  therein,  species  variations  be a b e t t e r c o n t r o l l e d  cellulose  changes  treatment,  f o r these  The t e c h n i q u e may  i.e.,  alpha-cellulose  structure of carbohydrate  variable.  1.  given  by  and  i t i s for holocellulose.  in  these  i n holocellulose  f o r a number  of gross  wood  a relationship  and c e l l  o f t h e age e f f e c t ,  length.  analyses  of the nine  s i n c e both  content  weight,  tracheid  increase with of years analyses  between  B u t , due t o  the relationship  age,  as d e s - " done  by  carbohydrate confounding  obtained  was  attributed obtained for  to  growth  factors.  Dadswell  a highly significant  radiata  pine,  suggested  et  correlation  a_l. ( 2 8 ) , (1 p e r  who  cent  level)  that:  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 can p o s s i b l y be cont 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 m o s t advantageous i f high average f i b e r length 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 .  Ifju ficant lose the  'r' value  content  addition,  ship. (3 8,  length  and  pine.  They  end  walls  result pulp  erial and  gross  cell  and  lignin  lignin that  (and  less  and  this  lignin.  diameter  are  in a  of  should  cellulose  and  they  results  less found  highly correlated,  the  et a l .  fibre loblolly in  fewer  consequently  compounds)  length  i n t u r n means  pulp  could -result  would  a  relation-  f o r s l a s h p i n e and  of  pectin  in  Einspahr  between  of  stem.  explain  content  studies of  fibres  Further,  single  age  length  unit  "longer  fibre  explained  per  content,  less  cellu-  " t h a t more  by  wood  wood  explanation  relationships  i . e . , increased  gram  gross  supported  given weight  less  per  wood  occur  also  signi-  increase with  content  an  a highly gross  authors  t r a c h e i d s , " which  negative  suggested  per  in  and  noted  was  and  basis of  offered  short  length  concept  yield";  fibres  of  These  the  lamellae w i l l  row  who  on  obtained  length  cellulose  authors  tracheid  This 39)  l e n g t h and  middle  also  tracheid  obtained  the  longitudinal observed  (60)  for Douglas-fir.  tracheid  compound  Kennedy  between  correlations  both In  and  and in  a  higher  fewer  intercellular that  from  fibre  which  mat-  length  they  51..  concluded may  also  that change  In gross lar  wood  stem,  ship  the  relative  due  view  to  of  the  holds  for different  and  whether  approach  can  be  at  30  noted  years  loblolly  pine  only  and  trees. was  slightly  lose  age  the  only  more  per The  work  length  one  layers  size.  of  a  similar  of  Zobel  correlation  a particu-  and  a  et  between  species. a l .  for  relation-  One  (17 2 ) . \  tracheid  length  a l p h a - c e l l u l o s e , among coefficient,  so  that  tracheid  per  cent  of  the  These  308  although  length  explained  variation  in  cellu-  content.  Even  though  i t i s apparent  tracheid  wood  carbohydrate  content  stems  of  species,  r e l a t i o n s h i p between  a  cause,  i s an  it  be  may  content  and  in  measured become authors  of  appreciated  since  cells  area  the  single the  a given for  less  relatively that  cell  former  volume  or  weight  cells. this  show m o r e  the of  than  species  and  its  between  cellulose  such  gross as  estimate wood,  per  the  such  cent  length,  of  and  e x p l a i n why two  between  Moreover,  Therefore, may  and  and  information.  characteristics,  s e n s i t i v e and to  little  on  length  related within  comparisons  i s based  individual  failed  are  that  gross  weak  wall  within  species,  correlation  two  cell  cell  to whether  of  cent  0.153,  than  in  characteristic  a between-tree  of  significant,  in  the  r e l a t i o n s h i p between  cell  as  stems  i t is  found  and  arises  of  increase  age-related  content  question  species,  authors  general  the  cellulose  proportion  are  numerous latter  is  comparisons some o f  the  variability  in  cellulose  content based  findings length  do p r o v i d e  to  dual as  results  9) d o s h o w t h a t ,  exists  f o ra strong  cell  both  obtained  irrespective  of species,  relationship  between  with  cause  wa's t o r e l a t e  from  are,highly  coniferous  weights,  When a c o n d i t i o n e d z e r o ) was f i t t e d  indivi-  However,  i s due t o chance, there  i.e.,  i s a common  between  species.  considered.  objective  of this  to i t s length  woods  from  (excluding  (see Figure  and  by t h e l e n g t h  model  those  from  data com-  are plotted.  relationships  f o r 91.9 a n d 95.7 p e r  i nholocellulose  regression  In  combined  trees)  the curvilinear  respectively,  investigation  i n conifers.  i s shown when  s i g n i f i c a n t and a c c o u n t e d  cent of the v a r i a b i l i t y  o f an  species  weight  obtained  (Figures',:l  skeleton.  and wood f o r m e d by f e r t i l i z e d  correlations  skeleton  be  10, a d e f i n i t e r e l a t i o n s h i p  wood  between  the possibility  length  a relationship  the primary  tracheid  a l l the nine  pression  age, o r whether  between  noted,  these  i ti s not apparent a t this  obtained  aspects w i l l  Variation As  workers,  f o rsuch  These and r e l a t e d  Figure  f o rr e l a t i o n s h i p  i n the present study  the relationship  increase  fundamental  2.  clues  Nevertheless,  c o n t e n t i n wood.  n o t e d b y some e a r l i e r  that  length.  and w e i g h t o f i t s carbohydrate  stage whether  The  earlier  and carbohydrate The  on c e l l  alpha-cellulose factor  alone.  (where t h e i n t e r c e p t i s  11), the variability  accounted f o r  was t h e same whereas"it  (91.9 p e r cent)  was s l i g h t l y  In vidual could  spite  species  less  o f t h e strong  characteristics  diameter  and w a l l t h i c k n e s s  a  relationship  pective of  slopes  these  of the species  (101, 102, 1 1 6 ) .  statistically  analyses  examined, respect to  Thus,  and w e i g h t  despite  i n each  f r o m one a n o t h e r ,  the regression lines.  i s given  i n Tables  and a l p h a - c e l l u l o s e t r a c h e i d  indi-  Such d i f f e r e n c e s  are also different with  and l e v e l s . o f  separate  holocellulose  for alpha-cellulose.  between l e n g t h  differed  weights,  relationships obtained,  s i n c e , f o r each  tracheid  some o f t h e s e  (95.2)  d i f f e r e n c e s were s i g n i f i c a n t .  be e x p e c t e d  strong  f o rholocellulose skeleton  species, i n res-  A summary  V and V I ,  for  skeletons,respec-  tively . Analyses V)  ..indicate that, i n spite  arity  i n regression line  family.  Thus P i n u s  menziesii, one  another  sitchensis sion  lines.  related of  interest  slopes  of the differences, there  slopes  lambertiana,  belonging  between species Picea  i n a d d i t i o n , Pinus  Taxus  not differ  t o note  the similarity  slopes  i n slopes  which  Cephalotaxus..  species  together  with  Taxus,  regres-  are closely  or levels. between  from  and P i c e a  o f t h e same  and 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  f o r the latter  t o one  and Pseudotsuga  lambertiana  and Cephalotaxus, i n either  i s simil-  d i d not d i f f e r  showed no d i f f e r e n c e i n t h e l e v e l s Similarly,  (Table  belonging  sitchensis  to the family Pinaceae,  i n slopes;  (159),did  dacrydiodes  and  f o rholocellulose tracheid skeletons  I t i s  Podocarpus  similarity i n Juniperus  The b e h a v i o u r with  some e x c e p t i o n s , i s  similarities related ties of  families  differ  lambertiana  from one a n o t h e r  i n slopes.  showed  significantly.in  either slopes  or levels.  Pseudotsuga d i d not d i f f e r  skeletons,  of regression  i td i f f e r e d  Pinus  and P i c e a ,  skeletons,  from  from Pinus lines  Possibly  this  relationship  matter  latewood.  Another  from  point  Araucaria,  the ratio  pattern  of  Therefore,  alphapossibly  may b e  dif-  i t would be o f i n t e r e s t to determine  and a l k a l i  f o rdifferent species,  riot-differ  i n the removal of  of shrinkage  further i n order  and  and Sequoia d i d  of the tracheids  between shrinkage  skeletons,  Picea  f o r holo-  those of h o l o c e l l u l o s e , could  i n different species.  to pursue  and  (5% l e v e l )  between  d i f f e r e n t behaviour  r e l a t e d to the shrinkage  hemicelluloses.  Interes-  as w e l l as t h a t between A r a u c a r i a  In addition, Araucaria  The s l i g h t l y  with  Juniperus, d i d  significantly  i n slopes  and  t h e same b e h a v i o u r with  levels.  fibre  Again,  two, together  Podocarpus.  ferent  Pinus  and Cephalotaxus  interest i s the similarity  cellulose  and  class.  the s i m i l a r i -  to different families, but  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 .  Sequoia,  be  Sometimes, however,  belonging  t h e same f a m i l y , i n s l o p e s  these  t o t h e same f a m i l y o r  and these  although  cellulose  in  Taxus  skeletons,  of h o l o c e l l u l o s e , i n showing  belonging  VI).  i n species  to slopes,  tingly,  and  that  sitchensis d i d not differ  regard  of  (Table  t h e same t r a c h e i d l e n g t h  Similarly,  of  like  between species  noted were  Picea  not  of alpha-cellulose tracheid  treatment  including  -  an  exact  i n these earlywood  3.  Variation with  The tracheid  strong  length  considered however,  and  individual weight,  together,  that  great  d i f f e r e n c e s can  this,  variation  was in  studied. Tables  has  f o r any  of  as  w e l l as  already  occur  age  species  and  f o r two  shown i n F i g u r e  12.  length  f r o m m a c e r a t e d wood o f  this is  butt  gravity values  log of  the  taken  that  increase about  150  f o r both  i n t r a c h e i d w e i g h t up years.  tracheids),  as  samples weighed  Tracheid  w e l l as (Tables  and to  length  the  I t i s known,  data  I I I and  and  i n the  years  listed  tracheid  tree  are  for tracheid  Warren  (70)  same f i g u r e . there  and  is a  tree  followed  and for It  marked  a decrease  after  (macerated  tracheid length  IV),  are  Radial  of  radius  c e r t a i n increments,  for this  average  the  500-year-old  latewood  80  property,  In view  across  length  from Kennedy  earlywood  species  physical  c o m p a r i s o n , mean v a l u e s  wedge s a m p l e , were a l s o i n c l u d e d  seen  or  Douglas-fir trees.  i n the  specific  out.  length and  weight patterns  obtained  f o r a l l the  w i t h i n a s i n g l e stem.  tracheid weight  For  that  anatomical  tracheid weight  IV  r e l a t i o n s h i p s between  been p o i n t e d  particular  Average  I I I and  increasing  the  of.the same  general  trend.  It sity to  pattern  the  study,  i s of  interest  described  by  to note  Kennedy  tracheid weight-length indicating  tracheid  that  and  and  radial  Warren  pattern obtained  some i n t e r - r e l a t i o n s h i p  characteristics  the  gross  (70)  (wood)  is  i n the  between  wood s p e c i f i c  den-  similar present  individual gravity.  The  56.  literature is  a v a i l a b l e , however,  possible in  that with  wall  from  i s comparatively  clearly  cell  increase  percentage of c e l l  which  It  does n o t show t h i s  (47). I n d i r e c t evidence  morphological  i n length  there  closely  greater  f o r longer  be an  I ti s  increase volume,  i.e.,  f o r longer  cell tracheids.  tracheid length i s  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  (138),  cell  fibres;  t h e r e f o r e , t o assume t h a t  Thus, van B u i j t e n e n pine,  may  w a l l as a p o r t i o n o f t o t a l  volume p e r u n i t l e n g t h would be h i g h e r  i s reasonable,  studies.  tree.  from h i s study  on s l a s h  concluded: T h e r e appear t o be two complexes o f p r o p e r ties which are rather strongly correlated. 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 summerwood 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 hand, 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 percentage, l o n g f i b e r s and a h i g h s p e c i f i c g r a v i t y on the o t h e r hand. These two c o m p l e x e s a r e t y p i c a l l y the j u v e n i l e p r o p e r t i e s versus t h e mature propert i e s , a l t h o u g h a p p a r e n t l y even 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 each other. This a s s o c i a t i o n , however, i s not 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 of combinations. One c a n , f o r i n s t a n c e , f i n d w o o d samples w i t h s m a l l f i b e r diameter, 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 , a n d a n a v e r a g e s p e c i f i c gravity.  The  reasons  specific to  gravity within this  the increase  tracheid heid  which,  length.  McElwee and  f o r a r e l a t i o n s h i p between t r a c h e i d length could  a l s o be a t t r i b u t e d  i n amount o f c a r b o h y d r a t e  fraction  as shown a b o v e ,  correlated with  i s highly  N o t much i s known o n t h i s  (171) f o u n d  specific  species  and  a poor  aspect.  i n the  Zobel  and  r e l a t i o n s h i p between c e l l u l o s e  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  trac-  strong  yield  57.  relationship specific the  on a volume b a s i s  i . e . , "a g i v e n  g r a v i t y wood g i v i n g h i g h e r  same v o l u m e o f l o w s p e c i f i c  indicate  highly  cellulose length  significant  c o n t e n t and f i b r e  and d e n s i t y  (99).  g r a v i t y -wood."  length  fibre  (28, 6 0 ) , as w e l l  i t was  noted  a growth  to  that  i s highly  ( r =,0.92) w i t h  possible  that  cellulose  specific  content  lose  specific gravity  properties  percentage,  i s thus  such  specific  of the i n d i v i d u a l tracheid  (Douglas-fir  level  (Figure this  (41).  found  12).  entirely  cellulose  Data  Increment  i n mature  Statistical  tracheids  from  this  cor-  I ti s to  and lower  cellu-  of latewood, show  skeletons  from  juvenile  and o v e r m a t u r e wood  indicate that  i t has  also  10), apparently,  analyses,  i s called  study  are well  and  that  below  increments  however, d i d not  (see Tables V I I and V I I I ) .  skeletons  the pith  g r a v i t y , percentage  weights  the  i s similar  significantly  t o m a t u r e wood  as-shorter  strength  1  increment  related indirectly  near  compared  tensile  wood  the pattern  g r a v i t y . (156, 1 5 7 ) .  i n the region  o r c o r e w o o d ; when  different  which  that  as  i n the tracheids.  Wood f o r m e d juvenile  studies wood  h o l o c e l l u l o s e v a r i a t i o n across  related  Several  relationship;', between gross  Also,  strength  high  y i e l d s of c e l l u l o s e than  of  of tensile  volume o f  Analyses  no s i g n i f i c a n t  confirm  f o r holo-  difference 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 a s j u v e n i l e w o o d , s i n c e M c K i m m y (94) s h o w e d 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 d o 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 a s j u v e n i l e w o o d .  58.  weight  could  be  found  compared  with  Contrary  to this,  (Table  VIII)  juvenile pared  wood  red that  fraction  i n wall From  these  could  also  loblolly from  7.5 wood  pointed  crown.  As  during  mature  The  conclusions  i . e . , f o r t h e same have more  hemi-  and overmature  wood.  qualitative  composition  with  increase  i n age f o r  t o be  c o n s i d e r a t i o n s , he inherent  Zobel  a n d may Some  and McElwee's  t h a t , based  per cent those  compared  concluded by  typified  indirect  (171) w o r k  on d r y weight,  higher  support on  yields  f o rw a t e r - r e s i s t a n t  f o r alpha-cellulose to the y i e l d  from  were  juvenile  trees.  o u t by L a r s o n  i s associated with  meristem,  exist;  com-  definite  found  when  i n weight  very  (WRC), w h e r e a s higher  length, the  (B) t r a c h e i d s .  i n the earlywood.  r a n 3.5  lower  conflicting  may  from  skeletons  tracheid  tracheids w i l l  and o t h e r  from  wood t r a c h e i d s .  some  appear  They  f r o m - t h e same  As wood  this  m a t u r e wood  per cent  those  chemical  found  pine.  wood  (78) h a s n o t e d  expressed  be  carbohydrate  f o r t h e same  differences  than  changes  the.trend  and overmature  apparently  juvenile  t r a c h e i d s as  f o r alpha-cellulose  that,  f o r these  length,  pine.  wood  (A) a n d o v e r m a t u r e w o o d  Larson changes  mature  analyses  that qualitative  cellulose  juvenile  tracheids are significantly  reason  tracheid  by  from  indicate  t o mature  obvious is  those  between  (76), formation  the prolonged  the growing  t h e c r o w n moves  season,  upward  of  influence of the i n the region  i n the older  juvenile apical  of the  tree,  the  active  cambium  a t a given  influences formed lower  height  of the elongating  (111). weight  Therefore,  evident,  since  wood  of  different quality  decrease  trees  (that  formed  higher  centage  than  Kennedy  and Warren  gravity  when  stresses dense  wood  formed  he e x p l a i n s  specific that  been  and s p e c i f i c  gravity i n  and V I I I ( C ) ;  i n the outer  lignin  portions)  and lower  during  thereby  earlier  stages  Doerner s  appreciably  stress  of growth. (3 5)  i n  decrease i n suggests bending  small  of  section  of v e r t i c a l  to  the tree  continues  stimulus  by p r o d u c i n g  the development  per-  the,formation  the rate  markedly,  thin-walled  i n height,  necessitating  when  over-  relatively  Doerner  ages,  to counteract  that  concept  1  a relatively  to the reduced  of  the  alpha-cellulose  t o augment  has decreased  be  Figure  has c h a r a c t e r i s t i c a l l y  i s growing  increment  and  (52) e x p l a i n e d  principles.  wood  period i s  quality will  on m e c h a n i s t i c  A t advanced  d e n s e wood  which  and subsequent  modulus.  efficiently  the juvenile  wood  to explain  and C l e r m o n t  (70) s u p p o r t e d  are greater,  peripheral  i n juvenile  the increase  the tree  significantly  offered  VII(C)  ( 3 0 0 t o 400 y e a r s )  prosenchyma.with  which  Hale  wood i s  cell.  have  (see Tables  F o r example,  m a t u r e wood old  above,  i n c e l l u l o s e percentage  o v e r m a t u r e wood 12).  arguments  f o r the  during  to the d i r e c t  and a d u l t  skeletons  produced  as s t a t e d  subject  region  the reason  i n the i n d i v i d u a l  Several  less  crown  of alpha-cellulose  is  reflected  becomes  to  radial react less-  o f an e x c e s s i v e l y  large  60. s e c t i o n modulus. changes to  For both  are produced  the stimuli.  age  effect  infers  independent  Results  of tree  one s i n c e  directly  diameter, whereas  diameter  tissues  support  a  direct  Doerner  i n c r e a s e s w i t h age.  on s i n g l e  tracheid  behaviour  of trees  o r r e a c t i o n wood  (Increments  No.  formation  are reflected  i n which  cell  (see below).  o b t a i n e d by t h e a n a l y s i s  (Increments  No.  Additional  of covariance  tracheid weights  differences  i n tracheid  length.  'slopes' i n both  cases  differ.  T h i s means  that  weights  i n overmature  300, 400), a f t e r  that  of over-  adjusting f o r  lower  c o u l d be a t t r i b u t e d  as t h i n w a l l e d prosenchyma c e l l s  wood  'F' v a l u e s  these populations  the s i g n i f i c a n t l y  wood  test  of mature  The s i g n i f i c a n t  indicate  more  whether i n  20, 8 0 , 150) w e r e c o m p a r e d w i t h t h o s e  m a t u r e wood  on  point of proof i n  i n the development of the i n d i v i d u a l  (Tables V I I and V I I I )  such  (52) t h u s  of the present study  i n growth  c o n f i r m a t i o n was  for  the density  of developing  a n a l y s e s , and p r o v i d e a f u r t h e r  changes  normal  authors  that  a r e i n agreement w i t h t h e above c o n c l u s i o n s based  g r o s s wood that  he s u g g e s t s  by t h e r e a c t i o n s  The f o r m e r  an i n d i r e c t  weights  cases  tracheid  t o some  as s u g g e s t e d  factor  by H a l e  and  Clermont (52).  4.  Variation  between r e g u l a r  compression  (normal)  and  wood  R e a c t i o n wood o f c o n i f e r s , p r e s s i o n wood, i s d e v e l o p e d  usually  referred  as p a r t o f an o r i e n t a t i o n  t o as  com-  mechanism,  i.e.,  a geotropic reaction  stated  t o an i n e r t i a l  force  (151).  As  by L a r s o n ( 8 0 ) :  The i n t e r a c t i o n b e t w e e n 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 thick cell walls. 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 the high auxin environment i n c l o s e p r o x i m i t y t o t h e crown. Chemically, the normal m e t a b o l i c pathways appear 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 duction of-constituents normally confined to the outer w a l l layers i s perpetuated across the wall.  Wardrop  and Dadswell  tracheid number the  length  eccentric  development similarity the  (Chapter  of a n t i c l i n a l  rapid  between  I n many made w i t h  pression  divisions radial  such  wood  of  may  i s usual  wood.  Thus  there  cells  of the  between (mildest similar  compression t o extreme to those  wood  normal  comparisons  wood.  o f same  the t o be  other  a  during  t h e main  frequen-  t o t h e comhowever,  length,  Moreover,  since  i n spite  difference  i s one o f d e g r e e  developmental  o c c u r r i n g i n n o r m a l wood  been  study,  tracheid  and s e n s i t i v e . differences,  have  adjacent  In this  a n d n o r m a l wood  type),  appears  with  tree.  the quadrant  morphological  with  and the t r a c h e i d s formed  investigations,  m a t e r i a l from  in"the  associated  that  be more m e a n i n g f u l  the obvious  an i n c r e a s e  growth  c o m p a r i s o n s a r e made o n t h e b a s i s they  from  the reduction i n  i n t h e cambium  of the l i f e  to typify  out that  II) results  of compression  youthful period  tly  (146) p o i n t e d  features  (see146).  only  being  Average and  their  mean  values  lengths  of compression  are contained  holo-  and a l p h a - c e l l u l o s e  ships  between  13.  tracheid  I t i s apparent  wood,  there  tracheid  i s again  weight  variation  a highly  and l e n g t h ,  compression  by  the length  wood  reaffirms  f o r n o r m a l wood  A normal  wood  significant  general  significant  factor  comparison  indicates  that,  difference  between  found  tracheid  skeletons  seem  to exist  for alpha-cellulose  existence  Examination the  original  length,  wood  differ  than  (where  In addition, from  wood  2.3 0 mm)  tracheids  mm).  however,  compression  under  13).  of the relationships data,  those  show  tracheid  weights, of  relation-  tracheid respect  wood a n d length,  differences  weights,  that,  f o r t h e same have  13 a s w e l l  slightly  tracheid  amount compared  suggesting  differences.  i n Figure  may  higher  lengths  were  t o mature  tracheids  i n holocellulose  as  tracheid  were m o s t l y , o v e r  wood  no  to holocellu-  given  lengths  wood  established  quantitative  compression  o f normal  cent  behaviour  In contrast,  wood (where  but a lower  since  This  skeleton  tracheids  juvenile  between  skeleton  compression  with  of q u a l i t a t i v e and/or  alpha-cellulose mainly  (Figure  compression  ,9) .  lose  the  alone.  f o r t h e same  can be  Figure  71.7 a n d 9 0 . 0 p e r  the previously  (see Figure  of  correlation  and a l p h a - c e l l u l o s e  for  Relation-  a r e shown i n  i n the formation  explaining  weights  I and I I ,  respectively.  and weight  even  i n holocellulose  respectively,  ship  that,  tracheid  i n Tables  skeletons,  length  wood  2.30  d i dnot  content,  they  must  contain  more h e m i c e l l u l o s e  wood  tracheids,i.e.,  fraction  the difference  than  between  comparable  normal  h o l o - and a l p h a -  cellulose .  The influence. sion  wood  data  I).  are  with  comparable  The Australian degrees form  only.  include to  bring  wood  here  (27).  may  amount o f  Since,  t h e extreme out these  to those  of  slightly juvenile  from  the mildest basic  cells,  wood  degree.  i t was  of  or i n the  and t h e d i f f e r e n c e c a r e was  of  that a l l  i n t h e mechanism  of daughter  of compression of  the  t h e r e i s no  wood,  wall,  tracheids  the findings  emphasized  be e n c o u n t e r e d  and normal  differences  length  principle.  with  i n the present study, case  from  alpha-cellulose  o n t h e same  t y p e s , and t h a t  of c e l l  Hence  Similarly,  authors  i n the readjustment  development  the pith  i n tracheid  behaviour.  are consistent  wood  t h e compres-.  f o r m a t i o n , t h e age e f f e c t i s  as compared  These  age-dependent  i t can be proposed  tracheids.  values  these  to  73 a n d 83 f r o m  a decrease  lower  wood  extreme  between  progressive  despite  be e x p l a i n e d  results  workers  division,  background,  slightly  could  be r e l a t e d  Increments  i n the cambial  normal  to the very  degree  this  o f compression  difference cell  from  alpha-cellulose  tracheids  could  i n the present instance,  compression  superimposed  higher wood  With  with  produced  than  was  obtained that,  associated s t i l l  behaviour  F o r example, selected  (Appendix the  above  i si n  taken to possible  In ship,  summary, i t c a n be s t a t e d  developed with  n o r m a l wood  that  tracheids,  the basic  relation  seems t o h o l d f o r  c o m p r e s s i o n wood  a s 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  anism i n c o n i f e r  cambial morphology.  wood  tracheids  differ  some q u a l i t i a t i v e drate  of the tree  emphasized  5.  from  species  difference  material  per tracheid  conifers  (14, 68, 121a).  specific  gravity  largely  to differences  substance  and  level,  formation.  latewood  that  earlywood increment,  the differences  zones w i t h i n  of a single tree  and  late-  have and  between  as, i fnot  a single  species,  evidence  i n the actual  i n earlywood  across  at cellular  be e s s e n t i a l l y as g r e a t  ( 5 9 ) . However, a n a t o m i c a l  significant  remarkable  factors, i s clearly  a growth  respect  t h o s e b e t w e e n wood  two members  carbohy  p h y s i c a l and chemical p r o p e r t i e s  I n almost every  than,  The  o f c o m p r e s s i o n wood  woods, w i t h i n  t w o t i s s u e t y p e s may  greater  i n showing  to the basic  readjustments  been r e c o g n i z e d  morphological,  compression  tracheids  regard  about by e x t r i n s i c  has long  of coniferous  behaviours.  wood  with  V a r i a t i o n between earlywood  different  these  f o r quick  i n the physiology  It  no  differences  changes brought  wood  f r o m n o r m a l wood  skeletal fraction (alpha-cellulose).  ability to  However,  mech-  stem,.  o r between  two  seems t o i n d i c a t e  amounts o f c e l l  and latewood  wall  tracheids i n  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 an i n c r e m e n t have been a t t r i b u t e d  i n t r a c h e i d volume r a t h e r  (70). I t i s of interest, therefore,  than  wall  t o examine t h e  variation  of individual  tracheids  fractions  of the increment. As a p r e l i m i n a r y  approach  length-weight relationships for  earlywood  relationships  and latewood. of best f i t ,  c o n i f e r o u s woods,  Y  t h e two  to this,  The c u r v i l i n e a r  ALPHA-CELLULOSE  Earlywood  Earlywood  0.0506X  = -0.3770 +  Y = -0.2418 +  2  R  0.198;  2  =  DF =  0.0310X  + R  2  DF  R  0.263;  =  0.0412X 0.936;  =  0.0301X  0.255;  R  2  =  DF =  X = tracheid  2  0.959; SEE =  weight length  +  0.0341X  Latewood 0.2214X 2  0.965; SEE =  0.152;  742  of tracheid i n mm  0.169;  383  +  Y = ovendry  0.2602X  Y = -0.3019 +  0.1201X 2  SEE =  0.130;  356  Earlywood  = 794 Where  2  Latewood  = -0.1708 +  2  0.972; SEE =  +  2  DF = +  0.0377X  0.1845X  Y = -0.3705 +  = 447 Earlywood  regression  Latewood  0.2365X  0.939; SEE =  Y  multiple  below.  Latewood  DF  separately  HOLOCELLULOSE  2 0.951; SEE = r = DF = 3 4 5  +  the tracheid  obtained are re-examined  +  Y  arbitrary-  f o r t h e combined data of t h e nine  are given  0.0291 +  =  from  i n g x  - green  10  condition.  These the  relationships  wood  within  correlation  confirm  that,  the increment,  between  length  irrespective  there  of origin  of  i s a high . degree o f  and weight  among  tracheids,  The  2 individual and  their  those  r  values  standard  between that, (six  of estimate,do  analyses  data  and latewood,  as w e l l  for holocullulose,  earlywood  species  examined  Interestingly, tests are  f o r t h e combined  always  heids,  confirming  ( 1 , 52,  significant their  114).  differences counterparts  Because tests  were  done  based  This  much  data, from  cases data  on  of  differences ' t 'tests  indicate  tracheids frequently higher  tendency  than  those  i s less  five  any s i g n i f i c a n t  pro-  of the nine  difference i n  fractions  (Table I I ) .  ( h o l o - and a l p h a - c e l l u l o s e ) , ' t ' showed  higher  that  latewood  i n weight  the findings  than  of others  I t i s t o be n o t e d , exist,  latewood  i n earlywood  o f t h e above  t o compare  holo-  the behaviour  s k e l e t o n s , where  t h e two t i s s u e  significantly  thus  analyses  than  i n both  o f both  o f latewood  I ) .  d i d n o t show between  done  further  Results  (Table  f o r alpha-cellulose  weight  not differ  are significantly  nounced  tracheid  and latewood  as t o d e t e r m i n e  weights  species)  tracheids  were  to explore  and w i t h i n s p e c i e s .  of the nine  f o r earlywood  values.  detailed  alpha-cellulose  earlywood  of  errors  o f t h e combined More  and  obtained,,  tracheid  earlywood based  however,  on  tracgross  that  where  tracheids are longer  (Tables  reason,  tracheids  I and I I ) .  analyses  weights  of  after  covariance adjusting for  length  differences.  Tables  IX  and  skeletons, the  no  mentioned, 'levels' the  nine  and  of  VI).  contrast  while  the  coniferous  within are  woods.  are  differences  species  significant  described  (i.e., only  are  in  found  between  IX).  As  expected,  which,  as  already  i n the,'slopes'  and  i n t e r e s t , however, i n that  latewood heavier  species.  such  differences  weights.  This  examined, but to  are  i s not  also  only  for a l l  o v e r a l l behaviour,  found between species  six Douglas-fir the  could  c h a r a c t e r i s t i c s of  given  significantly  regard  at  differences  of  between  trees In  X)  latewood,  a b o v e , no  six Douglas-fir  l a s t mentioned  ready  and  the  be  (Table  point  and  skeletons  to  can  are  holocellulose  differences  The  earlywood  tracheid  significant  ferences The  V  for  (Table  i n alpha-cellulose skeleton  for  highly  that,  examined  a t t r i b u t a b l e to  (Tables  analyses  differences  earlywood, both w i t h i n  true  these  seen  Douglas-fir  behaviour  evident  X)  of  are  In  the  be  of  e x i s t between species  holocellulose than  I t can  significant  s i x trees  differences  is  X.  Results  5 per well  cent  (Table  t r e e s ) , the level  dif-  (Table  r e s u l t from  the  j u v e n i l e wood and  IX).  a l -  compression  wood.  In tracheid it  summarizing  skeleton  i s evident  the  weights,  that,  f o r the  v a r i a t i o n s i n earlywood  f o r a l l the  same t r a c h e i d  l a t e w o o d w o u l d have s i m i l a r amounts o f differing  amounts o f  nine  hemicelluloses.  and  coniferous  length,  woods,  earlywood  alpha-cellulose, Qualitative  latewood  and  yet  differences  68.  in  the hemicellulose  workers but,  from  gross  i n addition,  cellulose tissue  with  In this Meier's  estimate for  as w e l l .  respect  also  been  analyses  Meier  adds  and  give  on  late-  only  on i n d i v i d u a l c e l l  weights.  to reflect  of individual tracheids  an  of carbohydrate  to the d i f f i c u l t y  a r e n o t s e n s i t i v e enough  f o r weights  based  about t h e  earlywood  analyses  (percentage)  which  however,  r e s u l t s a r e i n agreement  As n o t e d , . g r o s s fraction  other  i n alpha- .  (90) o b t a i n e d  f o rboth  by  ( 1 , 78, 90, 114)  I n one s t u d y ,  the present  based  reported  show d i f f e r e n c e s  pine,  as a whole,  data  analyses  values  reports  of alpha-cellulose  of the total  interpreting gross  these  findings.  the tissue  have  and t i s s u e  on Scotch  same p e r c e n t a g e wood.  wood  percentage  analysis  fraction  from  of  That i s , the true  different  origins.  Evidence  f o r a Common  From between  length  evident  that  conifers  in  of tracheid  length  origin.  i n the slopes  In other  As e x p e c t e d ,  considerations,  rather  than  lines  with  age  are s t a t i s t i c a l  depending  strictly  a common r e g r e s s i o n  a l l  one i n t h a t  increase there  for  i t i s  words, t h e  an age dependent  of regression  practical  fraction,  be d e r i v e d  and c e l l u l o s e c o n t e n t  a species.  I f ,from  statistical  on t h e r e l a t i o n s h i p s  and i t s carbohydrate  i s not necessarily  within  differences species.  observations  i r r e s p e c t i v e o f wood  tracheid  a stem  foregoing  i n Conifers  a common r e l a t i o n s h i p c o u l d  relationship both  Relationship  on  academic  line  could  be  fitted part  f o r a l l these  of the v a r i a b i l i t y  limits, zation ning  of standard  wood b a s i c  Thus, a g e n e r a l  The despite  conifers, tracheid  indicating  by  length  considered  estimates vidual  including  wood  a r e no h i g h e r (e.g.,  Alpha-cellulose), factor  with  entirely  gical  Figure  indicating  regard  that  done h e r e i n ,  would  wood.  since  mechanisms  over Thus,  within  a wide range of i f 'tracheid' shown i n  the correlations the standard  obtained  errors  of  f o r some o f t h e i n d i and F i g u r e  t h e paramount importance  improve  a  governing  a r e t h e same  9--Holocellulose  length-weight the effects  as c e l l  that,  different  the relationships  adding  c h a r a c t e r i s t i c s , such  10 c o n f i r m  tracheidsof  those noted  to the c e l l  possible  deduced.  between species,  formed  In addition, than  i s minimized.  of Figure  and w e i g h t r e l a t i o n s h i p s  significant.  be  the physiological  as one p o p u l a t i o n ,  species  could  among  10 a n d 1 1 a r e j u s t i f i e d ,  highly  introduced  factors  i n slopes  exists  that  randomi-  variability  relationships  relationship  a  of.the  and o n t o g e n e t i c  combined  Such  tolerable  a unique measure f o r exami-  age, n o r m a l wood and r e a c t i o n  Figures  not  since  significant  weight, within  behavioural pattern  between species,  tree  is  unit,  a  and c h e m i c a l b e h a v i o u r  s t a t i s t i c a l differences  significant  are  would provide  environmental  explain  of the estimate.  fundamental morphological  species,  is  i t would  i n tracheid  error  of tracheids  conifer  and  data,  of  1-length  relationships. of other  diameter.and w a l l  the relationship.  I t  morpholothickness,  As has been  70.  shown,  these  conifer the  cambial  more  based  three  in  any  of  wood.  morphology  dominant  on  the  morphological  one.  length  tracheid In  the  In length  and  direct  one,  on  the  length  in  that  is  difficult  of  cell  are  not  As  that of  in  the  the  to  pointed  elements  approach of  or  wall  of  the  the  same -  from  that  the  i t is  the is  f a c t o r s which  cell  a  amount  fraction  following  a  depends  indirect  influence  of  function  relationship is  an  - •  cell  finite . factor.  development  car-  "• - . - .  carbohydrate  some  that,  origin  r e l a t i o n s h i p between  whether  since  the  consideration,  hypothesize  to  is  be  carbohydrate  i r r e s p e c t i v e of  considerations  of  to  one, This  the  cell  extent  division  understood.  out  by  Bannan  i s determined  cambial  slight  in  Bayly  showed  that,  are  this  amount  known w h e t h e r  respond  and  relatively  ray.contacts  of  appearing  individual tracheid  amount  assess,  fully  to  an  cell;  inter-related in  factor  examined..  strong  a priori  variables  fusiform  (10)  a  i s not  enlargement  coniferous  woods  such  and  It  in  both  s t i l l  nating  of  fraction  i t s length.  the  are  s i m i l a r f o r .tracheids  i t i s reasonable  carbohydrate of  alone,  be  coniferous  view  advantage  estimated  would  weight,  morphology,  be  length  w o r d s ,• f o r . p r a c t i c a l -  bohydrate. f r a c t i o n •length.;, i n  One  factor  could  other  with  features  cells,  conifers. in  important  (8),  eventual  l a r g e l y by  size  size  of  of the  derived origi-  post-divisional elongation In  an  earlier  conifers, cell factors  for  size  being  work  Bannan  and  and  extent  of  s u r v i v a l of  cambial  daughter have  cells.  best  shortest  fail  or  feed-back  tant  initiation  age  of  provided  with  of  the  new lies  ray  rays in  with  tracheid  cell  length,  for  appears  the  from  tracheid  and  exist  consistent with  physiological  the  build-up  histories the  these  between length.  findings  mechanisms  derived a  effec-  equidisrays  growth,  contacts  of  of  and  to  well abun-  commensurate  cells  rigid  to  stor-  are  access  of  consideration the  amount  The  results  reported  governing  growth  with  framework  the  in  that  unit  length  than  sion  also  reached  longer  shorter earlier  determine . f i b r e  tracheids  ones by  of  as  by  a  direct  carbohydrate  obtained others,  herein  in  that  length-weight  according  relationship described  implication  that  (see  above have  has a  Figures  Clark  w£'i';ght p e r  (24), unit  to  10  who  size.  another  and used  length  important  weight  11), fibre of  the  predeter-  their  greater  are  relation-  relationships possibly  originating initials,  The  to  provide  an  initials  p o s s i b l y have ray  an  matures.  fraction  in  would  longest  of  the  t r a n s f e r and  sustenance,  the  balance  may  mined  where  proper  relationship  are  the  for  importance of  contacts  creates  provides  The  ray  whereas  This  function  they  nutrients  It  ships  (167).  for  best  develop,  which  contacts A  the  initials.  their  initials;  ray  nutrients.  ness'  new  with  and  r e g u l a t i v e system  dant  sufficient  cells  survive  nutrients, required of  thus  to  form  initials  division  the  longest  opportunity  tive  fusiform  The  a  per  conclu'coarse-  commercial  sulphite longer per  pulp  tracheids  unit  length  positively (108,  140)..  If  cations  i s so, such  A  on wood  applied  tracheids that  to test  compression  tion  to  raise  yield  to  to produce  o r wood  indicate  that  f o r shorter  which  have  undoubtedly,  tracheids.  important  was  between  wood.  impli-  t o have  apply  as e f f e c t s o f  land  since  compression  i s a  already wood  On may  morphogenetic  or reorientation  thing  of a soil  little  development  such  to  I t i s t o be noted, however,  regulatory  fertilization level  taken  The r e l a t i o n s h i p has  s u c h wood.  non-productive  appears  that,  thickness  of view.  i t i s a natural  the f e r t i l i t y  tilization  and w a l l  f o r longer  on t h e one hand,  f o r stem  as c h e m i c a l  or bring  activity  of normal  i n trees  automatically  differences  wood,  a n d , as s u c h ,  such  than  volume  i s correlated  of tips,  t o problems  properties.  and t h o s e  phenomenon  exists  wall  Relationship  r e l a t i o n s h i p as d e v e l o p e d  been  length  a r e l a t i o n s h i p may  of the  of cell  somewhat d i f f e r e n t a p p r o a c h  fertilizers  tor  i s more  point  As n o t e d , f o r  56, 158)  contribution  the p r a c t i c a l  Applicability  the  ( 5 0 , 54,  (146, p.391),  from  since  the p o s s i b i l i t y  the relative  wood.  percentage  be h i g h e r  diameter  Also,  less,  this  of coniferous  the average will  with  tracheids, weigh  samples  that  trees  the other  respond  hand,  be a r t i f i c i a l , i n order  into  d i r e c t e f f e c t on  t h e cambium c a n m o b i l i z e  i s no  a  fac-  created  to increase  production.  there  func-  Also,  the fer-  cambial  firm  evidence  nutrients  directly  73.  from  the  primary ment; the  transpiration or  and  d i r e c t e f f e c t on i t s e f f e c t on  r e s u l t of  following  the  processes  in  crown  the  see  that  what  is  to  in  a  are  above  happens  operative  crown,  wood  of  tree  the  (70).  and  viewpoint,  n o r m a l wood  (a)  might  be  length  be  (b)  an  of  increase  sibly  the  (18,  139);  decrease  no  change  tree  in  in  the  way  to  the  tracheid  spite  reverts  producing  best  fertilized  in  rate  to  a  of  of  per in  length-weight that  of  to  to  the  interest relationship  the  action  of  amount  since  there  surface  unit a  cells  period  possibilities  relationships obtained  for  pos-  due  of  to and  since  the  juvenility to  increase  cell divisions..  would  obtained the  and  period;  consequent cambial  will  foliage  length  given  per  increasing  overproduction  anticlinal  these  be  e f f i c i e n c y of  temporary  shorter  examine  t r e e s , .with  cells  that,  exist: "  photosynthate,  of  and  speak.  response  owing  c e l l weight  to  physiological  carbohydrate  photosynthetic  by  The  in  to  its  develop- •  i t appears  in photosynthetic  overproduction (c)  gain  of t r a c h e i d  availability  root  length-weight  in  possibilities  so  exerts  secondary  i t would  tracheid  a  on  f e r t i l i z e r , a l l the  Three  unit  Thus  activated,  the  also  i s both  (79).  fertilization.  There  Fertilization  growth  responses  addition  From to  stream  be  to  from  combined  compare the  data  for  conifers. present  For  case,  obvious  i . e . , because of  b e t w e e n some o f sons  can  reasons,  only  the  be  species  made w i t h  this  c a n n o t be  'slope'  studied. the  and  done i n  'level'  In view of  the  differences this,  r e l a t i o n s h i p obtained  comparifor  n o r m a l D o u g l a s - f i r wood.  Before riate  to  i n v e s t i g a t i n g these  t r a c e the  starting  changing  pattern  before  the  fertilizati6n  shown i n F i g u r e  14,  for a  was  fertilized  at  increment width, tracheids  per  It  seen  can  be  ages and  21  increment,  fertilization.  crease  ( o r one  due  to  the  their above. may  behaviour  i n weight  was  photosynthate  comparison,  to  to the  based  or  remained  three  after  of  the  years  on  holo-  four  data  after  temporary  macerated  same  figure.  cases  fertilization.  similar  to  the  i n latewood the  increased  cambial  general  again  after  dec-  year  of  a  de-  only which for  only  Subsequently,  tracheid weight  region,  this  by  The  pattern  availability  alpha-  thereafter  decrease,  of  for  immediately  c o m p r e s s i o n wood.  a  is  50  f o l l o w i n g the  followed  length,  which  and  static  refertilization);  i n weight of  both  year  increase  h a v e b e e n due  a pattern  latewood h o l o c e l l u l o s e skeletons,  f o l l o w i n g the  This  For  f o r two  formation  were the  showed i n c r e a s e year  year  In  a sudden i n c r e a s e  exceptions  one  decreased  i n weight continued  was  22.  Such  approp-  and  Douglas-fir tree  t h a t t r a c h e i d weight of  following  there  and  be  tracheid weight  period.  29-year^old  i t would  were a l s o p l o t t e d i n the  skeletons  application  of  mean t r a c h e i d l e n g t h  cellulose  rease  aspects,  described (year  23)  of  current  already  meeting  the  new  needle  requirements  (49,  80).  Further, this  an  increase i n the h e m i c e l l u l o s e f r a c t i o n  is  evident for alpha-cellulose Both  14)  mean t r a c h e i d  and  average  tracheid  (Tables  I I I and  IV)  for in  tracheid increment  width  fertilization further  tracheid  tracheid  length of  growth  apparent  changes they  o t h e r s , more f e r t i l i z e d  had  was  from  form  been s e l e c t e d ,  one  reasons  tree  each  response (Urea)  and  22,  16 of  (Figure weighed  to year s t i l l  noted  following 23  first  t h e r e was  quite  are  of  high  no as  changes  However,  are d i r e c t l y  of  increment  As  noted, Urea,  of  NPK„  and  and  another  (NPK,  they  this  study  Two  incre-  refertilization),  treatments were t h a t ,  i n increment width,  in  compositions  (NH^^SC^.  following  to  Douglas-fir,  treatments were s t u d i e d .  these  or decrease  from  three f e r t i l i z e r  (including these  cores  not  variation  related  of  in  i t is  independent  wood m a t e r i a l  of  namely  for selection  generated increase  and  year.  a steep increase  I n o r d e r t o examine the v a r i a b i l i t y  employed.  m e n t s , i . e . , 15  21  fertilization.  tree  (119),  tracheids  provides evidence  length or whether  i n the  i n that  fertilization.  length.  available  increase  increment  t h e r e was  r a t e was  tracheid and  primarily  same g e n e r a l p a t t e r n s  subsequent  following  the  a l l the  expected,  above p a t t e r n  weight  c l e a r whether in  but  to that before  The  length f o r the  f o r the years  ( y e a r 21);  increase,  compared  As  no  skeleton weights  f o l l o w e d the  weight.  since  was  showed  The while a l l  also  (NH.)„S0.) i n s p e c i f i c  definite gravity  following tracheid  fertilization properties  treatments.  f o r the three  The  m a i n p u r p o s e h e r e was  ton  weights per unit  observed (60)  showed a h i g h  percentage version length as  of c e l l  from  cell  as w e i g h t ,  specific already  are given  gravity.  Ifju  wall  which would  area  to density  cell  and Kennedy  Since  and w a l l  of relating  be e x p l o r e d .  gravity.and  a possible  values.  diameter  a possibility  g r a v i t y may  suggest  skele- -  t o changes  c o r r e l a t i o n between s p e c i f i c wall,  and  i n Table X I .  the carbohydrate  and t o r e l a t e these  specific  i s correlated with  well  trees  t o determine  length,  i n g r o s s wood  Some s e l e c t e d w o o d  concell  thickness  cell  weights  to  Such a r e l a t i o n s h i p has  been i n d i c a t e d f o r the 500-year-old  Douglas-fir  tree  (Figure 12).  Relationships  between  tracheid  length  and weight, i n 2  the to  fertilized 18, a l o n g  For  Douglas-fir  with  that  obtained  e f f e c t i v e comparison,  also  given  among  i n each  the treated  Table XII. (Table  figure.  are indicated i n Figures  f o r Douglas-fir  that  done i n each  _A s u m m a r y  of the overall  and u n t r e a t e d  Douglas-fir  no s i g n i f i c a n t  skeletons,  normal  the analyses  I t c a n be i n f e r r e d from  X I I A)  cellulose  trees,  trees  t h e combined  difference  b e t w e e n t h e n o r m a l wood  15  wood.  case are comparisons  i s shown i n analysis  exists, tracheid  f o r holoweight-  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 skeletons of the Urea t r e a t e d t r e e i s d i f f e r e n t from t h a t of normal D o u g l a s - f i r . Statistical analyses, however, i n d i c a t e t h a t the apparent d i f f e r e n c e s a r e n o t s i g n i f i cant. 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) of normal Douglas-fir (Figure 9 — H o l o c e l l u l o s e ) i s about three times t h a t o f t h e Urea t r e a t e d t r e e and thus i t would i n c l u d e a l l values of the l a t t e r .  length  r e l a t i o n s h i p and t h a t  t r e a t m e n t was (see  Figures  (NH N0 4  be  3  of f e r t i l i z e d  compared i n d i v i d u a l l y  with  wood.  normal  15 t o 1 8 ; H o l o c e l l u l o s e ) , w i t h  treatment—Figure  Douglas-fir  one  1 8 ) , no s i g n i f i c a n t  each  exception  difference  could  observed.  Contrary alpha-cellulose (Table  X I I A)  dually  with  t o t h e above,  skeletons  among  same b e h a v i o u r was  and w i t h i n n o t e d when  skeletons' from  either  or level  out significant  the f i v e  difference that  i n slopes,  the behaviour  respect But  skeletons,  (Figures  i s , they  Urea  they  d i d not differ  these  treatments  differ  Thus,  i f the regression  length,  than  may  XII B),f o r significant This  implies with  values,  was  between slopes.  i t c a n be c o n c l u d e d  that  e f f e c t s o n wood  of best Urea  holocellulose tracheids  comparable ones  exception  i s different  i n either levels or  i n their  equations  f o r predicted  i n that  i s no  treatments  The o n l y  differences,  values  there  (Figure 15).  (Table  are parallel.  i n these  In view o f the l e v e l  examined  Interestingly,  length-weight r e l a t i o n s h i p , i s similar.  differed i n levels.  a n d NPK w h i c h  indivi-  of the d i f f e r e n t treatments,  the magnitude of response  since  The  treatment Urea d i d not d i f f e r  indicate that  (response)  to tracheid  trees.  16 t o 1 8 ) .  from normal D o u g l a s - f i r  that  differences  Douglas-fir  Comparisons between the treatments holocellulose  analysis f o r  treatments were compared  normal Douglas-fir  i n slope  t h e combined  points  alpha-cellulose  are  When  properties.  f i t (Figures  15 t o 18)  t r e a t m e n t shows  higher  are heavier  fromthe other  per unit  treatments.  I ti s  78.  therefore  possible  are  responsible,  the  observed  fertilized  lose  changes  skeletons.  their  with  lines  respective difference  the  XI),  for  i n these  XII B).  when  With  regard  S(  those  of the other  skeletons  f o r the r e l a t i o n s h i p  Although  t h e above  length  relationships  tional  proof  f o r the thesis  governing  comments  again  t h e same  for  increased  are taking  rate  means  In this  the  individual  to tracheid  they  also  i n that  that  while  place,  weight-  provide  addi-  the physiological relationships  i n the f e r t i l i z e d  this  weight),  confirming  weight-length  b i o l o g i c a l viewpoint,  of  gravity.  wood,  the  growth  thus  refer  hypothesis  the tracheid  and  are similar to  between  specific  i n fertilized  tracheid  treatments.  for holocellulose,  a n d g r o s s , wood  NPK  15 t o 1 8 ) , t h e U r e a  (predicted  f o r alpha-cellulose  Urea  a r e made o n t h e b a s i s  f i t (Figures  values  i n  to levels,  )  comparisons  higher  (NH^)  (NH^) 2 - 4 k u t n o t f r o m  of best  reported  NPK  a n d NH^NO^ d i d n o t s h o w  results  i s once  of holocellu-  Urea  while  than  weight  that  dif-  slopes,  showed  tracheid  than  from  and  again,  advanced  skeletons  Urea,  (Table  argument  conifers  specific gravity  length,  findings  mechanism  (see Table  of treatments  equations  tree  respect,  weight  e x i s t between t h e  Here  unit  wood  cell  Significant differences  NH^NO-j.  per  factors  i s somewhat d i f f e r e n t  s i g n i f i c a n t l y from  treated  with  of alpha-cellulose  differed  regression  associated  other  i n gross  behaviour  treatments  regression  such  along  changes  trees.  The ferent  that  trees.  i n  From  readjustments  the suggested  relationship temporary weight While the  i smaintained.  reversal  to juvenility,  among  r e l a t i o n s h i p there  the treated  trees  i ntheir  for  effects of fertilizers  properties with a  studies  growth  tree has this  temporary clinal lated  processes  ( 8 ) . I n view  (Table  XII B).  ficant  differences  holocellulose those  difference this  years  cells.  regression  lines,  exists  tracheids  with  associated  I ti sevident  that  of reverting to rate  of anti-  These a s p e c t s  and r e -  extensive  works o f  o f j u v e n i l e wood  i ti sclear that  e i t h e r i n slopes  tracheids  U r e a , NPK a n d ( N H ^ ^ S O ^ . ( i nslopes from  o n l y ) when  But a  comparing  t r e a t m e n t NH^NO^ .  This  be a t t r i b u t e d t o t h e age f a c t o r  t h e NH^NO^ f e r t i l i z e d  of first  no s i g n i -  or levels of  b e t w e e n " j u v e n i l e wood  those  could  F o r example,  o l d a t t h e time  length  wood a n d t h a t  analyses,  c a n be found  i nbehaviour  tree.  i ntracheid  o n wood  e f f e c t s , c o m p a r i s o n s w e r e made  of fertilized  From t h e s e  differ-  properties.  as a r e s u l t o f an i n c r e a s e d  difference  j u v e n i l e wood  o n wood  type o f mechanism  from t h e treatments  significant  thelimits of  indicating the possibility  (71, 108, 117).  of such  length.  magnitude o f response  a r ew e l l documented in:the  between behaviour  of  rate  remarkable  juvenility  i ntracheid  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  d i v i s i o n s i n thecambial  Bannan  and  of tracheid)  have shown a d e c r e a s e  increased  i n tracheid  w e r e some s t a t i s t i c a l  length  Many  changes  case, w i t h i n  (weight p e r u n i t differential  i s achieved by a  o f changes  seems t o b e t h e g e n e r a l  suggested  this  so that  arebasically a reflection this  ences  Obviously  fertilization  t r e e was 21  (Figure 1 4 ) ,  80. whereas to  t h e U r e a , N P K , (NH^)  juvenile period—see  holocellulose from  those  tracheid  of juvenile  Analyses differences tracheids  point,  NPK.  treatment  line  from  two t r a c h e i d  those  populations  l i n e s between  n  o  The i m p l i c a t i o n s  i n slopes  o f these  carbohydrate  in  i nlevels of  skeletons  S 0  4  (Figures  t h e amount o f a l p h a - c e l l u l o s e The b e h a v i o u r  length  of tracheid.  16 a n d 1 7 ) d i f f e r e d perunit  length  skeletons  NH^NO^ i s s o m e w h a t d i f f e r e n t f r o m t h e o t h e r s . normal D o u g l a s - f i r ,  i n that  S0  normal Douglas-fir,  of tracheid  regression  are important  only  15) d i d n o t show a n y s i g n i f i c a n t d i f f e r e n c e , 2  from  (NH^)2 4 w i t h r e s p e c t t o  fraction perunit  compared w i t h  (NH^)  Urea and from  or levels of  findings  resulting  b o t h NPK a n d  4  The i n t e r e s t i n g  show s i g n i f i c a n t d i f f e r e n c e  U r e a a n d NPK d i f f e r e d f r o m  F u r t h e r m o r e , when  S 0 2  suggesting  and treatments  alpha-cellulose t  wood  (NH^)  U r e a , NPK, a n d  differ.  j u v e n i l e wood  to this,  thejuvenile  o f j u v e n i l e wood,  treatment  (Figure  between  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  t h o s e o f j u v e n i l e wood e i t h e r lines.  slopes  i n d i c a t e no s i g n i f i c a n t  i sthesignificant difference  (NH^) 2^4  Consequently  wood.  Once a g a i n  In contrast  close  t r e a t m e n t NH^NO^ d i f f e r e d  f o r alpha-cellulose  i n slope  however,  regression  from  a g e 13 ( v e r y  fertilized.  and those f o r t h e treatments  significantly these  were  X I ) when  skeletons  i nregression  (Table X I I B ) .  that  Table  trees  t h e NH.NO-. t r e a t e d  tree  Urea  whereas significantly  of tracheid.  from  treatment  When c o m p a r e d t o (Figure  18) s h o w e d  81. significant cellulose from the  differences  skeletons,  these curve  two t r e e s i n Figure  cates  that  these  unit  length  the  18  that  ship  and  the tracheid  alpha-  populations  I n t e r e s t i n g l y , the upper p a r t  (Holocellulose—where may  X = 3.92)  of  indi- .  have more h o l o c e l l u l o s e p e r  c o m p a r a b l e n o r m a l wood  tracheids.  be a t t r i b u t e d t o t h e i n c r e a s e  conclusion,  suggested  maintained quickly  Such  i n weight  (years  from  to the increased  possibility per  unit  The  period  23 a n d  as e x e m p l i f i e d  o f temporary  as w e l l  and  age.  the  period this  growth  noted i n  24-—Figure  Where  the trees  of r a p i d growth  such  (46a),  i t may  the  expected  norm i n t r a c h e i d  the f e r t i l i z e d trees  reversal  and  (NH^^SO^.  and t h e d e g r e e o f  reduction  of increase i n  as i n i t i a l  tree  take  i n  speedily  I f the trees are  longer  before  length-weight.  they  In the  used were young,  and i t i s n o t known w h e t h e r  vigor  and s t i l l  they might recover  reversal to juvenility.  period,  adjust  (alpha-cellulose)  are growing vigorously  this  trees  In addition, the  b y t r e a t m e n t s NPK  f a c t o r s such  and p a s t  fertilization,  i n weight  relation-  and weight, i s  may w e l l b e r e l a t e d t o r a t e  old  instance  length  tracheids.  juvenility  as t o o t h e r  temporary  the basic  r a t e by a t e m p o r a r y  e x i s t s f o r a decrease  length  that  f e r t i l i z e d trees;  and produce s h o r t e r  tracheid length,  growth  i t c a n be s a i d  f o r normal wood, between  i n t h e wood  juvenility  from  holo-  following refertilization.  In  in  differ.  f o r both  latewood h o l o c e l l u l o s e skeletons  14),  to  suggesting  tracheids  than  b e h a v i o u r may  i n slopes  reach  present  a t the time of  the reported  trends  82.  in  t r a c h e i d weight  will  the  the  in  p r o p e r t i e s even i n o l d e r t r e e s ,  applied  i n moderation  silvicultural  and  Some P r a c t i c a l  Two tion, the  obtained  same f o r o l d e r s t a n d s .  theless, fibre  results  be  and  aspects  of  areas of  the  the  ethanolamine  o f wood and  been suggested  with  of  3050 t o  major polysaccharides yields  obtained  alpha-cellulose by  others  (20,  123,  are  slightly  ences  discussed  science  NaBH  t o be (163)  is  other  93  present I I ) , are  per  by  the  present  been  intermediate TAPPI  (chlorine-  peracetic acid  cent  recovery  f o r both  comparable  method  holocellulose  c a r b o x y l groups  study  and  industry.  that of  a superior  minimum o f  i n the  and  obtained  the  sec-  fraction  r e d u c t i o n ) , has  4  and  t o 100  in this  relationship  somewhat  However,  to give  to  of  (83).  holothose  and reported  135). t o be  i n t h a t the  higher  in yield  and  appears  acid)  be  fraction  (and  (83)  3600,  (Appendix  There (peracetic  change  (79).  carbohydrate  pulp  holocellulose  a d o p t e d has  The  of  holocellulose  (128)) m e t h o d s .  a D.P.  drastic  when f e r t i l i z e r  tracheid length-weight  originator  between c h l o r i t e  will  yield  method, u s i n g p e r a c e t i c a c i d by  any  consideration to  management p r a c t i c e s  practical  Yield  reported  indicate  Considerations  implication to  not  w i t h due  namely, experimental  obtained,  do  Never-  than  some p e c u l i a r i t y holocellulose  those  from  yields  latewood.  have a l s o been"recorded  by  for this for  Similar  Leopold  for  method earlywood differloblolly  83.  pine  wood  since as  (83) .  the  cooking  compared  seem  to  to  reason  cycles  five  influence  expected dix  The  for  the  for  latewood.  difference i n that  on  assess  yield,  such  have  such,  have  minor  addition,  will  earlywood But  latewood  the  this  is  obvious  were  only  behaviour  y i e l d s , which  values  based  and  However,  checks  amounts  of  of  some  the  can  be  tance  tracheid  and  vegetative  and  paper  viduals  within  to  three  does  not  showed  the  were h i g h e r  on  since  single  to  the  the  of  actual than  (Appen-  for  species  in  on to  As  will  be  adjusted see  for  that  a l l the  indicated  the  study,.it  the woods  comparable  work. the  re-emphasizing  fibre  Its  the  to  and  paper  impor- . in  the  s e l e c t i o n of  f i b r e s , not only  morphology  that  skeletons  individuals  importance  judicious longer  appears  carbohydrate  s e l e c t i o n of  having  r e l a t i o n s h i p between  obtained  based  employed  present  p r o f i t a b l y , by  lies  variabilities.  taken  samples  weights.  d i f f e r e n t woods  values  was  to  II).  the  propagation  that  tracheids,  difficult  tracheid  those  were  pulp  i t is  inherent  r e l a t i o n s h i p of  length  individual  assume  conditions of  used  industry  in  of  t h i s work, c a r e  light  obtained  made  due  (Appendix  length-weight  of  been  realistic  in  lignin  In  not  weight  incorrect  same d e l i g n i f i c a t i o n Spot  the  same y i e l d ,  comparisons  values.  to  differences  i t is  more m e a n i n g f u l  the  needed  difference  II) .  based  the  this  alpha-cellulose  Corrections  In  for  breedi pulp  indibecause  of  properties  84.  (13,  29,  34,  150),  carbohydrate length  and  increase tionate It  in unit increase  therefore pulp  weight  or  less to  yield.  weight  higher  its  amount  the  that  due of  this  of  Horn  This  per to  grow  or  trees  pulp  and  trees  concept  of  possessing  a  of  pulping  10,  would  11).  in  areas  a  per  given  result  point  in  related  per  gram,  and  c h a r a c t e r i s t i c s as  process.  have  every  dispropor-  result  fibres  (pulp)  for  (Figures  contact  been  fibres  volume longer  of  The  implica-  discussed  recently  oversimplification, i t  result  wood,  fibres  fibres,  industry;  of  (84)  i t would  rather  i f the  briefly,  i n more  than  wood  this  managed  optimum fibre  for  the  for  paper  quality  justified  seem  propagating to  be  used  yield  some y e a r s  production  and/or on  content,  advantageous  pulp  these  species in  the  forestry ago  f o r e s t s would  continuous  can  cellulose  a high  suggested  properties is  is  is  ' c e l l u l o s e f o r e s t r y , ' whereby  for  be  important  number the  risk  Similarly, Leslie  scientifically  Breeding  or  shorter  paper  will  could  wall  an  in  (58).  with  with  or  fibres  r e l a t e d aspects  i f longer  weight  concept.  yield  that  consequently  the  analyzing  considerable  that  given  on  meaning  cellulose  up  differences  r e l a t i o n s h i p between  one,  end  brings  based  Coens  At stated  and  fewer  possible  there  of  wood, w h i c h  species,  and  to  fibre,  longer  of  the  curvilinear  in  significance for  tions  addition,  of  evaluation  by  be  a  because  length  (39) .  affected  by  is  yield  volume  also  In  appears  lignin  pulp  but  of  the  be wood  making.  grounds.  85.  Moreover, nature  considerable  of f i b r e  Although  length  estimates  any  significantly  can  s t i l l  with  fibre  length,  shortening  pressure  as b r o u g h t  fitably, mining tion or  point  t o produce  higher  i s highly  may  i n d i v i d u a l company  i n p a r t i c u l a r f o r those striving  demands  point  of the pulp  of cellulose  length  since the  significantly  further because  fibre  correlated  consideration, of  increasing  returns.  The  c a n be employed  requirements, may  countries  to produce  d o n o t show  study.  merit  maximum  This  yields  for fibre  relationship of tracheids  t h e r o t a t i o n age.  increasing  c e l l u l o s e content  r o t a t i o n age o f t h e s t a n d  to suit  countries  wood  out i n this  that  as t o t h e h e r i t a b l e  37, 47, 61, 100, 154, 169, 1 7 0 ) .  by b r e e d i n g  composition  f o r land  length-weight  exists  h e r i t a b l e nature,  Another is  ( 8 , 9,  f o r gross  be o b t a i n e d  individual  evidence  be worth  s e t t i n g up  e n o u g h wood  and paper  i n deterinvestigaplantations  t o meet t h e  industry.  pro-  CHAPTER  RECOMMENDATIONS  M u c h .of weight  (carbohydrate  represents exists  a  was  and  and to  a  wide  applicable further  to  relationship mental  actual  A  this  results  of  are  practical  the  more  indicates  subsequent has  that  cell  wall  further  material  has  with  phenomenon  encoun-  and  been  more  a  weight  and  already  study  of  to  the  intra-increSince  difference  be  of  be  needing  tracheid  i t would  to  will  consideration.  per  regard  detailed  designed  conclusions  significant  tracheids,  primary  example,  further  that  Its  many q u e s t i o n s  For  no  relationships,  striking  so  raised.  tracheid  (stem) woods.  project  material  on  overall relationship  for  the  paper  in  the  between interest  tracheid  to  weights.  b e e n made h e r e i n ,  but  inconclusive.  relationship length,  this  individual fibre  approach  Other  and  an  RESEARCH  length  i n general,  latewood  aspect  preliminary  of  gravity.merits  evidence  and  of  been  between  amounts  earlywood pursue  have  specific  anatomical  Although  conifers  study  and  some b a s i s  range  in  coniferous  determine  investigations. include  skeleton)  within  provide  FURTHER  information  reconnaissance  between  objective tered,  the  FOR  V  studies  that for  and  may  that  need  further  e x i s t between  angiosperms,  biological  so  as  research  include  individual fibre to  significance.  evaluate  the  weights  results  for  both  87.  It in  tracheid  subsequent cellulose test  occurs  tracheid  i n this  A  degree  study  more r e a l i s t i c  i n developmental  cells,  accompanying  studies  could  parenchyma  ted  several  employed used for  This  be done.  of research  profitably.  studies,  o n wood  facilitates  strength same  density  direct  into  dimensions  to  can be  made'with could  elements  such  and Wellwood  single  be  changes  xylem.  fibre  as  vessels  (118) s u g g e s could  be  weights  can be  or strength/density  ratios  walls  studies  i n cell  wall  i n v o l v i n g wood and  The uniqueness  comparison  holo-  would  the chemical  the e f f e c t s of s i l v i c a l  properties.  This  tracheid weights  of cell  i n quantitative  properties, fibre.  where  strength-weight  and a s s e s s i n g  variables  Sastry  F o r example,  conducted  information  involving  differentiation  individual fibres,  tion,  the  areas  i n determining  porosity  thus  could  i n the  i s related to  comparisons  F i n a l l y , - w e i g h ing«o.f:other c e l l and  be  skeletons  r e l a t i o n s h i p f o r alpha-  of holocellulose skeletons.•  useful  reduction  fraction  and s p e c i e s .  length-weight  a  tracheid  of reduction  (earlywood/latewood)  so t h a t  that  i n alpha-cellulose  skeletons.  exact  study  of the hemicellulose  the rate'or  a more  cellulose, those  observed  t o removal  type  provide  in  length  whether  tissue  has been  forma-  environmental  of this  approach  o f c h a r a c t e r i s t i c s such  and c a r b o h y d r a t e  content  on  as  CHAPTER  VI  CONCLUSION  1.  A quartz  accurate  weighing  range  0.06  was  of  used  of single  t o 14  ug  and  cells.  The  woods,  been  a p r e c i s i o n ±0.03  i n individual  belonging  has  balance  f o r quantitative determination  alpha-cellulose ferous  ultra-microbalance  has  yg.  o f both  different  a  holo-  genera  balance and  of nine and  for  weighing  This  tracheid skeletons  to nine  devised  coni-  seven  families.  2. drate  Length  i n i t are highly  variations  accounted  alpha-cellulose, 91.9  and  95.7  physiological its  weight  by  Between  in  diameter  statistical  coniferous  the length  and  factor  between  skeleton)  species levels  and w a l l  alone,  The  i s  that  of  A  direct, and  suggested.  may  g r o u p e d as o n e  observed  lines.  have  For practical be  estimated  and  of tracheid  d i f f e r e n c e s were  thickness  could  length  carbohy-  were r e s p e c t i v e l y  studied.  of regression  behaviour.  tracheids  correlated.  for a l lspecies  relationship  to slopes  the amount.of  for i n holocellulose fraction  per cent  respect cell  t r a c h e i d and  significantly  (carbohydrate  3.  this  of a  with  Differences  contributed  to  considerations, population.  the  Small  2 standard  d e v i a t i o n , - and  tracheid  weight  and  highly  length,  significant  support  this  r  values,  argument.  between  89.  4. followed  The r a d i a l  an i n c r e a s i n g  subsequently  juvenile  cellulose  F o r t h e same wood  than  differences  trend  those  were n o n s i g n i f i c a n t  carbohydrate  amount t h a n  others:-  length. based  same to  wood  tracheid  length,  alpha-cellulose.  tracheids normal) for  may  wood,  t h e same  8. sical, of  tained both lose, being  i n this  earlywood  species study  heavier.  decreased  individual  tracheids  amounts o f  alpha-  wood,  whereas  for holocellulose.  had s i g n i f i c a n t l y  f r o m m a t u r e wood,  lower  f o r t h e same  to those  reported  tracheids  d i d not d i f f e r  amount o f h o l o c e l l u l o s e ,  but differed significantly  lower  concluded  that  alpha-cellulose  amount when  by  f o rthe  in,respect  compression than  compared  from  mature  wood (and  to juvenile  wood,  length.  properties have shows  and latewood  but differing  and  and overmature  Significant differences  -and c h e m i c a l  coniferous  wood  in.the  but higher tracheid  weight  analyses.  I t was  contain  lower  a r e comparable  Compression tracheids  150 y e a r s  length,  tracheids  those  Results  on gross  7. normal  tracheid  wood  tracheid  wood.  from mature  Overmature  of single  t o about  had s i g n i f i c a n t l y  6.  tracheid  up  i n the overmature  5. of  variation  long that,  i n morphological,  between been  earlywood  recognized.  f o r t h e same  had s i m i l a r  and  latewood  Evidence  tracheid  amounts o f  amounts o f h o l o c e l l u l o s e ,  phy-  ob-  length,  alpha-cellu-  latewood  tracheids  90.  9. tilized growth in  trees rate  have  also  part,  contribute  were  between there  length  weight, related  length  others. possible gravity  less This cause  formerly  i n addition  of carbohydrate  associated  exists  Evidence  compared  indicate  alpha-cellulose difference f o r changes  of fertilized  that  tracheid  Thus  per unit  i n behaviour observed  trees.  gravity.  f o r t h e most  wood;  wood  where  trees  of a  compar-  may,  however,  compared  suggested  i n gross  relation  juvenility,  when  may  Changes'  the basic  treatments  was  obtained  variables,  i n fertilized  length  i n  skeletons  t o n o r m a l wood some  was  t o t h e above  t o a phase o f temporary  tracheids  i n fer-  to differences  fertilization,  length."  s t i l l  o f wood  i n specific  with  to tracheid  and w e i g h t  shorter  gravity  of latewood.  t o an a l t e r a t i o n  age.' 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T a p p i 41: 167-170.  of  cellu-  172.  , T h o r b . j . o r n s e n , 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: 149-158.  173.  , G o g g a n s , J . F . , M a k i , T.E. a n d F. H e n s o n . 1961. Some e f f e c t s o f f e r t i l i z e r s o n w o o d p r o p e r t i e s of 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 SINGLE  TRACHEID  (HOLOCELLULOSE SKELETON)  SPECIES Araucaria Sequoia Pinus  cunninghamii  sempervirens  lambertiana  Podocarpus  dacrydioides  NO. OF OBSERVATIONS 46 40 35 28  Picea  sitchensis  25  Taxus  brevifolia  27  Juniperus  virginiana  Cephalotaxus  wilsoniana  33 21 204  Pseudotsuga  menziesii  Pseudotsuga  menziesii  Pseudotsuga  menziesii  116  Pseudotsuga  . . . menziesii  53  Pseudotsuga  menziesii  52  Pseudotsuga  menziesii  69  2  4  Combined  Average  48  797  I  WEIGHTS  TRACHEID DESIGNATION EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW  9,  AND  LENGTHS  FOR  NINE  TRACHEID LENGTH mm, if ic ± 1 SD 8.12 8.05 6.15 6.73 6.48 6.15 4.31 4.97 4.57 4.54 2.10 2.51 2.20 2.62 1.95 2.18 4.35 4.89 2.65 3.03 2.92 3.26 2.64 2.71 2.24 2.45 2.57 2.71 3.72 4.18  + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +  0.73 0.79 0.50 0.64 0.71 1. 29 0.64 0.57 0.91 0.90 0.32 0.30 0.47 0.36 0.17 0.20 1.65 1.52 0.38 0.36 0.56 0.56 0.22 0.27 0.32 0.27 0.24 0.37 1.82 1.88  CONIFEROUS  WOODS  T R A C H E I D WEIGHT g x IO x ± 1 SD -  3.43 3.54 1.66 2. 23 2.53 2.46 1. 40 1.72 1.08 1.08 0. 26 0.32 0.19 0.26 0. 21 0. 28 1.07 1.70 0.41 0.66 0 . 50 0.79 0.42 0.53 0.29 0.40 0.32 0. 47 0.90 1.27  + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +  (Continued)  6  0.75 N.S. 0.82 * 0.42 0.47 0.41 N.S. 0.86 0.33 * 0.29 0.42 N.S. 0.40 * 0.08 0.06 * 0.08 0.06 0.04 * 0.05 ** 0.67 0.90 ** 0.10 0.18 ** 0.22 0.30 0.14 ** 0.17 ** 0.08 0.11 ** 0.11 0.13 0.89 ** 1.06  106.  TABLE  a  A l l weights  EW  Earlywood  LW  Latewood  x SD N.S.  *  Average  Earlywood 5%level  (Continued)  expressed - i n ovendry  length  Standard  I  or  condition  weight  deviation and l a t e w o o d ('t' t e s t )  not  significantly  different  at  Earlywirod and l a t e w o o d 5%. l e v e l ( 1 ' t e s t )  significantly  different  at  the  E a r l y w o o d and l a t e w o o d 1% l e v e l ('t' t e s t )  significantly  different  at  the  the  1  **  Groups  within  Douglas-fir  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, 3 0 0 , 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.  Compression  3.  NH4NO3 f e r t i l i z e d  wood  (Increments and 22.  21,  - Breast height disk  of  a  20,  120-year-old  150  tree,  - Breast height disk of a 29-year-old tree 22, 2 3 , 24, 25) f e r t i l i z e d a t a g e s 21  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 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) 13 a n d 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 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 13 a n d 15.  6.  80,  (NH4)2SO4 f e r t i l i z e d - 12 mm i n c r e m e n t h e i g h t , 24-year-old t r e e (Increments a t a g e s 13 a n d 15.  from b r e a s t h e i g h t ; f e r t i l i z e d a t ages  height; a t ages  core from b r e a s t 1 5 , 16) fertilized  TABLE I I SINGLE TRACHEID  (ALPHA-CELLULOSE SKELETON) W E I G H T S NO. OF OBSERVATIONS  Araucaria Sequoia Pinus  cunninghamii  sempervirens  lambertiana  Podocarpus  dacrydioides  33 32 35 43  Picea  sitchensis  46  Taxus  brevifolia  27  Juniperus  virginiana  Cephalotaxus  wilsoniana  26 21  Pseudotsuga  menziesii  199  Pseudotsuga  menziesii  43  Pseudotsuga menziesii ; 4 Pseudotsuga Pseudotsuga  menziesii menziesii  Pseudotsuga  menziesii  Combined  Average  112 42 42 46 745  TRACHEID DESIGNATION EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW • LW EW LW  3  AND  LENGTHS  FOR N I N E C O N I F E R O U S  TRACHEID LENGTH mm, x ± 1 SD 7.38 7.35 5.36 5.81 5.37 5.81 4.17 4.07 3.98 3.95 1.93 2.13 1.87 2.12 1.41 1.64 3.33 3.88 2.31 2.62 2.16 2.70 2.08 2.39 1.99 2.18 1.97 2.33 3.12 3.56  + + + + + + + + + + + + + + + + + + + + + + + + + + + + +  0.78 0.80 0.81 0.60 0 . 62 0.39 0.48 0.45 0.51 0.46 0.39 0.29 0.24 0.27 0.32 0.27 1.21 1.27 0. 54 0.46 0.37 0.54 0.17 0.23 0.30 0 .25 0.31 0.46 1.55 1.57  WOODS  TRACHEID_WEIGHT q x 1 0 ~ 6 x ± 1 SD 3.20 3.07 1.58 1.81 1.92 2.30 1.19 1.14 0.89 0.88 0.18 0. 23 0.19 0. 23 0.14 0.16 0.95 1.26 0.40 0.52 0.33 0.62 0.31 0.43 0.29 0.35 0.24 0.37 0.79 1.01  + + + + + + + + + + + + + + + + + + + + + + + + .+ + + + + +  0. 53 0.73 0.48 0. 42 0.37 0.19 0. 24 0. 29 0. 24 0.19 0.06 0 .06 0.05 0.04 0.03 0.04 0. 63 0.74 0.17 0.17 0.13 0. 24 0.06 0.09 0 .07 0.08 0.09 0.12 0.77 0. 84  (Continued)  N.S.  ** ** N.S . N.S . N.S.  * N.S .  ** * ** ** * ** **  108. TABLE  a  A l l weights  EW  Earlywood  LW  Latewood  x SD N.S.  * **  Average  expressed  length or  Standard  II  (Continued)  i n ovendry  condition  weight  deviation  Earlywood 5% l e v e l  and latewood ("f test)  not significantly  Earlywood 5%level  and latewood ('t' t e s t )  significantly  different  at the  Earlywood and latewood 1% l e v e l ( ' t ' t e s t )  significantly  different  at the  Groups w i t h i n  different  at the  Douglas-fir  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 1 0 , 2 0 , 8 0 , 150 3 0 0 , 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.  Compression  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 ( I n c r e m e n t s 2 1 , 2 2 , 2 3 , 2 4 , 25) f e r t i l i z e d a t a g e s 2 1 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 1 5 , 16) f e r t i l i z e d a t a g e s 13 a n d 1 5 .  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 1 5 , 16) f e r t i l i z e d a t a g e s 13 a n d 1 5 .  6.  wood  - Breast height disk  (NH )2SC>4 f e r t i l i z e d - 12 mm i n c r e m e n t h e i g h t , 24-year-old tree (Increments a t a g e s 13 a n d 1 5 . 4  of a 120-year-old  tree, tree  core from b r e a s t 1 5 , 16) f e r t i l i z e d  TABLE I I I SINGLE  T R E E NO.  TRACHEID  (HOLOCELLULOSE SKELETON) W E I G H T S  INCREMENT NO. FROM P I T H  NO. OF OBSERVATIONS  10  27  20  26  80  23  150  39  300  26  400  27  20 21 2 2  a b c Tree Tree  36 23  b  b  24  23  28  24  15  25°  26  A l l weights expressed Years fertilized, Compression wood  i n ovendry  a  3  LENGTHS  TRACHEID DESIGNATION  2.17 2.41 3.58 3.98 5.61 5.85 5.97 6.26 5.79 5.46 5.85 6.00 2.92 3.20 2.92 3.40 2.93 3.22 2.98 3.34 3.34 3.58 2.67 2.86 x SD EW LW  condition  of a  FOR  TWO  DOUGLAS-FIR  TRACHEID LENGTH mm, x ± 1 SD  EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW ' LW EW LW EW LW  1 - Butt l o g of a 500-year-old tree; 2 - NH N0 f e r t i l i z e d - Breast height disk 4  AND  + + + + + + + + + + + + + + + + + + + + + + + +  0.30 0.38 0.70 0.66 0.70 0.52 0.60 0.48 0.69 0.95 0.61 0.71 0.59 0.48 0.57 0.33 0.45 0.54 0.48 0.72 0.23 0.27 0.73 0.52  TREES  TRACHEID_WEIGHT g x 1 0 ~ x ± 1 SD 6  0.24 0.39 0.85 1.26 1.65 2.25 1.77 2.62 1.63 1.90 1.42 1.87 0.56 0.84 0. 53 0.70 0. 54 0.74 0.44 0. 85 0.71 1.02 0.37 0.62  + + + + + + + + + + + + + + + + + + + + + + + +  Average length or weight Standard deviation Earlywood Latewood•  29-year-old  tree.  0.07 0.13 0.35 0.44 0.52 0.51 0.47 0.68 0.45 0.56 0.28 0.49 0.21 0.33 0.16 0.32 0.18 0. 21 0.12 0.36 0.28 0.32 0. 21 0. 20  TABLE SINGLE  TRACHEID  (ALPHA-CELLULOSE SKELETON) W E I G H T S  INCREMENT NO. FROM P I T H  T R E E -NO.  NO. OF OBSERVATIONS  10  28  20  34  80  24  150  27  300  25  400  26 35  20 21  b  22  22  b  20  23  26  24  22  25  a b c  22  C  A l l weights expressed Years fertilized Compression wood  Tree Tree  IV  i n ovendry  3  3  LENGTHS  TRACHEID DESIGNATION  condition?  FOR  + + + + + + + + + + + + + + + + + + + + + + + •2.54 +  1.85 2.18 2.94 3.46 4.22 4. 87 4.26 5.07 4.-01 4.76 4.48 4 .54 2.39 2.80 2.38 2.79 2.35 2.66 2.03 2.60 2.12 2.98 1.90  x SD EW LW  :  of a  TWO  DOUGLAS-FIR  TRACHEID LENGTH mm, x ± .1 SD  EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW  1 - Butt l o g of a 500-year-old tree. 2 - NH N0 f e r t i l i z e d - Breast height disk 4  AND  0.37 0.33 0.54 0.62 1.04 0.74 1.22 1.01 0.73 0.86 0.70 0.94 0. 42 0.42 0.44 0.47 0.26 0.28 0.19 0.71 0.33 0. 57 0.40 0.50  g  TREES  TRACHEID_WEIGHT x IO x ± 1 SD -  0.22 0.31 0.73 0.96 1.3 8 1.86 1.50 1.97 1. 24 1.79 1.33 1. 63 0.45 0. 66 0.36 0.64 0.32 0. 60 0.31 0.56 0.38 0.80 0. 27 0.51  6  + + + + + + + + + + + + + + + + + + + . + + + +  Average length or weight Standard deviation Earlywood Latewood  29-year-old  tree.  0.09 0.12 0.27 0.32 0.56 0.47 0.70 0.60 0.37 0.51 0.44 0.58 0.13 0. 21 0.15 0.22 0.08 0.17 0.07 0.28 0.15 0. 27 0.16 0.18  TABLE  V  A N A L Y S I S MODEL (A) AND (B) R E S U L T S OF 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 (HOLOCELLULOSE SKELETONS) ( A D J U S T E D FOR 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 ) A)  Analysis  Model  . Group  Differences Differences  DF  F  Total for testing slopes Sums  X  for testing levels Combined Regression Group  B) 1 vs 2 vs 3 vs 1 vs 2 vs 3 vs 6 vs 7 vs 8 1 vs 2 vs 4 3 vs 5 vs 9 1, 2, 4, v s 6, 7, 8 v s ! 1 vs 2 1 vs 3 1 vs 4 1 vs 5 6 vs 7 6 vs 8 6, 8 v s 4 vs 9 3 vs 5 1 vs 9  4 vs 5  Significance i n Slopes Levels  ** ** ** **  5 vs 9  ** ** ** * ** ** ** * **  N.S , * N.S , **  **  **  N.S.  ** *  *  **  N.S, **  N.S .  N.S , N.S , **  N.S .  **  **  **  Groups 1 2 3 4 5 6 7 8 9  Araucaria cunninghamii Sequoia sempervirens Pinus lambertiana . Podocarpus dacrydioides Picea sitchensis Taxus b r e v i f o l i a Juniperus v i r g i n i a n a Cephalotaxus wilsoniana Pseudotsuga menziesii  X  - Indicates 'Significance' or 'Non-significance' in slopes, Indicates 'Significance' or -'Non-significance i n levels 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 1  N.S. *  **  112. TABLE  VI  A N A L Y S I S MODEL (A) AND (B) R E S U L T S OF 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 (ALPHA-CELLULOSE SKELETONS) ( A D J U S T E D FOR 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 ) A)  Analysis  Model Group  Differences Differences  Total for testing Sums  slopes  f o r testing levels Combined R e g r e s s i o n  B)  Group 1 vs 2 vs 3 vs 4 vs 1 vs 2 vs 3 vs 5 6 vs 7 vs 8 vs 9 1 vs 2 vs 4 3 vs 5 vs 9 1, 2, 4 v s 9 6, 7, 8 v s 9 1 vs 2 1 vs 3 1 vs 4 1 vs 5 6 vs 7 6 vs 8 6, 8 v s 7 4 vs 9 3 vs 5 1 vs 9  DF  F  -  X  -  Y  Significance i n Slopes Levels  **  5 vs 9  N.S .  **  N.S.  * ** **  N.S . N.S . N.S . N.S.  *  N.S . N.S .  **  N.S .  **  ** ** ** * ** ** ** N.S .  **  N.S.  ** * **  N.S. •  ** ** **  Groups 1 2 3 4 5 6 7 8 9  Araucaria cunninghamii Sequoia sempervirens Pinus lambertiana Podocarpus dacrydioides Picea sitchensis Taxus b r e v i f o l i a Juniperus v i r g i n i a n a Cephalotaxus wilsoniana Pseudotsuga menziesii  X  - Indicates 'Significance' or 'Non-significance' i n slopes  Y  - Indicates 'Significance' or 'Non-significance' i n levels  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  113. TABLE 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 FOR T R A C H E I D WEIGHTS ( 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 , MATURE AND OVERMATURE WOOD OF 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 LENGTH) Group  A  DF  F  J u v e n i l e wood ( I n c r e m e n t " 1 0 ) M a t u r e w o o d ( I n c r e m e n t s 2 0 , 8 0 , 150) Total:  24 85 109  Differences  2 111  1 .190  N . S.  1  2 .010  N. S.  f o r testing  slopes Sums:  Difference Combined  for testing  levels  Regression  112  Group  B  DF  J u v e n i l e w o o d ( 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 300, Differences Differences Combined  for testing for testing  400) Total:  24 50 74 2 76  0 .722  N .s .  Sums:  1  0 .355  N. s .  slopes levels  77  Regression Group  C  F  DF  F  M a t u r e w o o d ( I n c r e m e n t s 2 0 , 8 0 , 150) O v e r m a t u r e w o o d ( I n c r e m e n t s 3 0 0 , 400) Total:  85 50 135  Differences  2 137  7 .401**  1  29 . 7 5 2 * *  f o r testing  slopes Sums:  Differences  f o r testing  levels  Combined R e g r e s s i o n ——— — — — — — — — — — — — ——— — —————————— — ———— — — N.S. **  Not s i g n i f i c a n t Significant  a t t h e 5%  a t t h e 1%  level  level  138  114. TABLE  VIII  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 FOR T R A C H E I D WEIGHTS ( 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 , MATURE AND OVERMATURE WOOD OF 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 FOR 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 ) Group  A  F  DF  J u v e n i l e w o o d ( I n c r e m e n t 10) 25 M a t u r e w o o d ( I n c r e m e n t s 2 0 , 8 0 , 150) 82 T o t a l : .107 Difference  f o r testing  slopes Sums:  Difference Combined  f o r testing  levels  Regression  B  400) Total:  slopes Sums :  Difference Combined  for testing  levels  Regression  7 .787  25 48 73 2 75  - 1 .413  1  15 .069  N. S .  **  F  82 48 130  Difference  for testing  slopes  2 Sums : 132  Difference  f o r testing  levels  1  N.S. * **  **  F  DF  M a t u r e w o o d ( I n c r e m e n t s 2 0 , 8 0 , 150) O v e r m a t u r e w o o d ( I n c r e m e n t s 3 0 0 , 400) Total:  Combined  N. S.  76  Group  C  1  DF  J u v e n i l e w o o d ( 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, for testing  2 .575  110  Group  Difference  2 109  Regression  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  133  3 .898  *  0 .034 N. S .  115.  TABLE IX A N A L Y S I S OF C O V A R I A N C E AND A D J U S T E D MEAN V A L U E S OF EARLYWOOD AND LATEWOOD T R A C H E I D W E I G H T S I N S I X DOUGLAS-FIR TREES ( 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 LENGTH). a  HOLOCELLULOSE SKELETONS Source  DF  Trees  5  Earlywood/Latewood  1  Error  ALPHA-CELLULOSE SKELETONS  F  DF  1.23 N . S . 97.92  **  534  F  5  2.45*  1  0.00 N.S,  475  Adjusted  Mean  Tracheid  Skeleton  Weights  Earlywood  0.7 5  0.70  Latewood  0.94  0.7 0  —6 a N.S. * **  A l l weights Not s i g n i f i c Significant Significant  e x p r e s s e d a s g x 10 a n t a t t h e 5% l e v e l a t t h e 5% l e v e l a t t h e 1% l e v e l  i n ovendry  condition  116.  TABLE  X  A N A L Y S I S OF C O V A R I A N C E AND A D J U S T E D MEAN V A L U E S OF EARLYWOOD AND LATEWOOD T R A C H E I D W E I G H T S I N N I N E b CONIFEROUS S P E C I E S ( A D J U S T E D FOR D I F F E R E N C E S I N T R A C H E I D LENGTH) a  HOLOCELLULOSE SKELETONS  ALPHA-CELLULOSE SKELETONS  Source  DF  Groups  13  20.91 **  13  1  65.43 **  1  Earlywood/Latewood Error  DF  781  64.66 ** 1.88  729  Adjusted  Mean  Tracheid  Skeleton  Weights  Earlywood  1.02  0.90  Latewood  1.17  0.91  a b,c N.S. **  A l l weights Includes a l Not s i g n i f i c Significant  N.S  e x p r e s s e d a s g x 10 i n ovendry c o n d i t i o n l species and t h e s i xtrees w i t h i n Douglas-fir; a n t a t t h e 5% l e v e l a t t h e 1% l e v e l  TABLE S E L E C T E D WOOD AND  TRACHEID PROPERTIES  Tree  Characteristics Treatment  First  fertilized  THREE  FERTILIZED  84  Tree  12 0.414  TREES  Tree  90  15* 0.430  12 0.403  16 0.436  74  (NH ) SO 4  15* 0.374  16 0.352  12 0.381  2  15* 0.371  4 16 0. 337  28.6 4.2 2.26 27.0  23.1 6.5 3.30 28.0  20.0 5.5 2.80 27.5  24.7 5.1 2.10 29 . 8  18.6 10.5 2.36 29.5  18.6 10.7 2.55 29 .4  24. 6 5.7 2.20 29.2  14.3 7.7 2. 26 27. 8  10. 7 7. 5 2. 40 26. 2  34.5  43.1  40.0  33.0  38.0  40.5  38.0  40.5  46. 0  7.6 9.0  a t the  DOUGLAS-FIR  NPK  age o f  7.5 10.0  13  7.0  P  c o n t e n t method.  7.5  10.5  o  r  K  7.5 12.0  11.0  and r e f e r t i l i z e d  of 200 l b Cpounds per acre of N, 2 ^ § ' 1. Maximum m o i s t u r e  OF  Urea  Composition  Increment from p i t h Specific gravityl ( 3 - y e a r age segment) Percentage o f latewood I n c r e m e n t w i d t h , mm Tracheid length, mm Tangential tracheid diameter, micron Radial tracheid diameter, micron Tangential double ,-;.wall t h i c k n e s s , m i c r o n Radial double w a l l thickness,micron  *  XI  2^ 3 i e <  u  v a  a t age  ^-  e n t s  )  7.5 12.5  15 a t  7.0  7.5  8. 0  11.5  10. 2  10. 5  the rate  118. TABLE X I I R E S U L T S OF 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 ANALYSES; 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 UNTREATED DOUGLAS-FIR ( A D J U S T E D FOR 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 )  A.  (HOLOCELLULOSE SKELETONS)  Group  DF  a  1 3 4 5 6 Total Differences Differences Combined  f o r testing slopes Sums f o r testing levels  Regression  B.  4 4 4 3 4 5 6 3  vs vs vs vs vs vs vs vs  5 vs 6 6 5 4 7 7 7 7  N.S. * **  a  DF  F  201 113 50 49 66  196 109 39 39 42  479  425  8 487 4  8 433 N .S . 4  1.115  **  7 .405  **  Significance i n Slopes Levels  ** ** **  ** ** N.S .  *  N.S N.S N.S N.S  **  5 .650  437  491  N.S. N.S. N.S . N.S . N.S . N.S . N.S .  F  N .S.  1.915  Significance i n Slopes Levels  Group  (ALPHA-CELLULOSE SKELETONS)  N.S . N.S . N.S. N.S.  . . . .  **  ** ** N.S . 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  Group  2 - Compression heading  wood  of Douglas-fir  is  under separate  (Continued)  119.  TABLE  Groups  within  XII  (Continued)  Douglas-fir  1.  Butt log of 150, 300,  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 1 0 , 20, 80, 400) 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 . ?  3.  NH4NO3 f e r t i l i z e d  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 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) 13 a n d 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 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 13 a n d 15.  - B r e a s t height disk of a 29-year-old t r e e (Increments,.'21, 2 2 , 2 3 , 24, 25) f e r t i l i z e d a t a g e s 21 a n d 22. from b r e a s t height; f e r t i l i z e d a t ages  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 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, a t a g e s 13 a n d 15.  7.  Juvenile  wood  -  Increment  10  from  Group  1  height; a t ages  from b r e a s t 16) fertilized  above.  120.  FOREWORD TO  Figures  1 t o 10, 13,  Mathematical  model  Y  15  t o 1 8 - - P l o t t i n g and c u r v e by c o m p u t e r .  f o r curve  = b  Q  FIGURES  + b  ±  fitting  fitting:  X + b  X  2  2  _g Where  Figure  In of  Y  =  Ovendry  X  =  Tracheid  weight length  of tracheid i n g x i n mm  - green  10  condition  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 . of b e s t f i t c o n d i t i o n e d t o pass through  a l l figures, best f i t .  the mathematical  models  given  The l i n e the origin,  are the  equations  Figure  1:  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) 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 samples from A r a u c a r i a cunninghamii.  H 3.0  I 0.98  I 1.96  I 2.94  1 3.92  1 4.90  Tracheid Length, mm'  1 5.98  1 6.86  1 7.B4  1 8.92  1 9 BC  Figure  2:  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) 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 samples from Sequoia sempervirens.  122, H0LO-SP2  A.  -r22.2633=+ 0, .6580KS = 0.3OS; DF = 38 0 , 6 6 4 W H E R EIY = O V E N D R Y W E I G H T OF TRACHEID N G . I O " X-=GRE TE RN ACHEIDCONDIL NN GTH IN ffl TE IO E  6  x  l'.96  —I 2.94  -1  4.9©  3.92 .  5.88  6.86  ALPHA SP2  -• 0.2 1590=+0.9O .; OSStXS = 0.119; DF = 30 3 3 W H E R EIY = O V E N D R Y W E I G H T OF TRACHEID N G . 1 0 6 X-=GRE TE RN ACHEIDCONDIT LI EO NN GTH IN HI 2  B,  E  x  —1 3.98  -r 2.94  3.92  4.90  Tracheid Length,mm-  J.flfl  Figure  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) 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 samples from Pinus lambertiana.  123. H0L0-SP3  A.  Yr.= -1.120 78 +S 0.6222= X 0.183; DF = 33 2 = . 9 3 3 ; W H E R EIY = O V E N D R Y W E I G H T CF TRACHEID N G . 1 0 6 X-=GRE TE RN ACHEIDCONDIT LI EO NN GTH IN W E  x  88  —I  0.90  —I 2.54  —1  —1 4.90  3.92  —1 E.B6  -1  ~1 8.82  T.B4  RLPHfi SP3  B.  -r1. 0880=+ 0; .5722XS = 0.130; DF = 33 2 0 . 8 6 0 W H E R EIY = O V E N D R Y W E I G H T CF TRACHEID N G . I O " X-=GRE TE RN ACHEIDCONDIT LI EO NN GTH IN Wi E  6  x  2.94  - r - — 3.92  ~T 4.9<?  Tracheid Length, mm-  ?.8e  Figure  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) 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 samples from Podocarpus dacrydioides.  124 H0L0-SP4  . 0 119=0+0.9290 .0t97S X = 0.109; DF = 26 'H 2E .YrW = R EIY = O V E N D R Y W E I G H T OF TRACHEID N G . X I f f * X-=GRE TE RN ACHEIDCONDIT LI EO NN GTH IN HI 2  E  (£>  0-.0  I 2.94  - 1 — 0.96  I 3.92  -1 5.88  -1 6.86  —1 7.64  RLPHfi SP4  B.  Yr2==0.01 0 1 +S0.05= 27X0.068; DF = 11 0 . 9 0 0 ; W H E R E : INYG. =O E N D R Y E I G H T OF TRACHEID xV 10-6W 2  E  X = TRACHEID LENGTH IN MM - GREEN CONDITION  I  1.96  T 2.94  —I 9.92  -r  4.90  Tracheid Length, mm-  - 1 — 7.84  —1 8.82  Figure  5:  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) 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 samples from Picea. s i t c h e n s i s .  H0L0-SP5  Y = -0.7389 + 0.H939X r = 0.948 2  S  EE  = 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  -1 3.92  2.94  —1 4.90  r  5.6  ALPHA SP5  Y = -0.0056 + 0, 1-2 = 0.920; S  EE  = 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  3.32  -i 4.9C  Tracheid Length, mm  r 5.B  -1  6.86  ?  6.82  Figure  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) 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 samples from Taxus brevifolia.  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  3.92  1  4.9«  Tracheid Length,mm-  —I  6.96  Figure  7:  Relationship between t r a c h e i d weight (holocellulose (A) a n d alpha-cellulose (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 from Juniperus v i r g i n i a n a .  127. HGL0-SP7  Yr2==0,01 1 0 +S0.03=47X20.025; DF = 31 0 . 8 9 4 ; W H E R EG Y =IOO V E N D R YW E I G H T OF TRACHEID ] . " X-=GRE TE RN ACHEIDCONDIL EO NN GTH IN P M TI E  6  x  —I  -1  1.98  -1  3.92  5.BB  —I G.B6  -1  T.B4  flLPHfi SP7  =r2-0 .10220.91+ 0.1 549X= 0.013; DF = 24 = 8 ; S W H E R EIY = O V E N D R Y W E I G H T OF TRACHEID N G . 1 0 6 X-=GRE TE RK ACHEIDCONDIT L E N GTH IN W I O N E  x  2.94  —I  3.92  4.90  Tracheid Length, mm  ?  a.82  9.Bp  Figure  8:  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) 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 ) a n d l e n g t h f o r s a m p l e s from Cephalotaxus w i l s o n i a n a .  128 . H0L0-SP8  A.  Y = -0.2790 + 0.2526X r  2  = 0.921; S  EE  = 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  1.96  -1  2.94  3.92  RLPHfi SP8  B.  Y = 0,0618 + 0.035GX  2  r2  = 0.818; S  EE  = 0.013; DF = 19  WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G, x 10-6 X = TRACHEID LENGTH IN FN - GREEN CONDITION  8  ^ 3.98  I 1.66  I 2.94  I 3.92  I 4.90  Tracheid Length,mm-  I 5.88  1 6.96  1 7.84  1 8.82  1 Q.80  Figure  9:  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) 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 samples from Pseudotsuga m e n z i e s i i .  129.  384  A. y = 00769+00554x  320  o  Condil  r  ,endi  >.  3  2  =0-843  2  EE =0-342  DF=202  2-56  Where y = oven dry weight of  o  tracheid in g xlO'  i  1-92  oX  x = tracheid length in mm -green condition  cn  a>  1-28  ochei  13  H  least-squares fitted line 0-64  0-00"— 000  4-48  0-98  1-96  2-94  y = -0-3189+0-2184 x +00437 x  3-92 490 Tracheid Length, mm  5-88  2  R =0-986 2  B.  B 3-84  S  o o  DF = I96  fT 3 20  T3  E E  =0-083  Where y = oven dry weight of tracheid in g xlO""  6  O  2 56  x= tracheid length in mm -green condition  •92  _ a>  least-squares fitted line  1-28 -  u  £ 0-64 000 000  0-98  2-94 3-92 4-90 5-88 Tracheid Length,mm-  6-86  7-84  6-86  7-84  8-82  Figure  10:  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) 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 from nine coniferous s p e c i e s . (Leastsquares f i t t e d line).  130 HOLO SP1-9  Y = -0.1463 + 0.099OX + 0.0436X  2  fi = 0.919; SEE = 0.319; DF =456 2  WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G, X 10"6 X = TRACHEID LENGTH IN MM - GREEN CONDITION  Tracheid Length, mm  Figure  11:  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) 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 ) a n d l e n g t h f o r s a m p l e s from nine coniferous s p e c i e s . (The l i n e of best f i t c o n d i t i o n e d to pass through the origin).  131 HOLD SP1-9  -R0. 03E8X0,91+ O .OW X0,320; DF = 457 = 9 , S = W H E R EY =.O V E N D R Y W E I G H T CF TRACHEID I N G x 1 0 6 X-=GRE TE RN ACHEIDCONDIT LI EO NN GTH IN ffl 2  2  E  a.a  1  1  r  0 98  1.96  i.94  1 39i  1 4.9o  1  5.88  r B.I  —I— T.B4  ALPHA SP1-9  =R0. 0758X0.95 + O. OW iX0.196; DF=459 = 2 ; S > W E R EIY = O V E N D R Y '-.'EIGHT OF TRACHEID N G , x 1 0 E X-=GRE T RN A.CHEID L E N GTH IN m E CONDIT I O N 2  2  E  -r  2.94  3.92  —r  4.9C  Tracheid Length,mm-  —i  8.82  Figure  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 across the b u t t l o g of a 500-year^old Douglasf i r tree. (Each p o i n t r e p r e s e n t s t h e average t r a c h e i d weight or length f o r the increment. Specific gravity values from Kennedy and Warren (70)) .  •_£T  A  Figure  13:  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) 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 compress i o n wood a n d n o r m a l wood o f D o u g l a s - f i r .  133 HOLO COMP WOOD  A.  Yr=2=-0.0 5. 97 51 97;+S0. 1012X0.103 = = 16 10-6  MULTIPLE CURVILINEAR COVARIANCE ANALYSES  21 05 1 212 6 2118 0.287N.S. 219  CROUP  DE  DOUGLAS-FIR (NORMAL WOOD) COMPRESSION WOOD TOTAL DIFFERENCE FOR TESTING SLOPES SUMS DIFFERENCE FOR TESTING LEVELS  WHERE Y = OVENDRY WEIGHT OF TRACHEID  IN  G. x  X = TRACHEID LENGTH IN MM - GREEN CONDITION  0.C60N.S,  COMBINED REGRESSION N.S,  E  DF  NOT SIGNIFICANT AT THE 5? LEVEL  , least-squares fitted line ^ f o r Douglas-fir  least-squares fitted line for Douglas-fir Compression Wood  ° s  —I 2.94  RLPHP, C O M P WOOD  Yv==0.00 21 1 +S0.C= 0.057 D E . 9 0 0 ; 11 90 6 DF = 11 232 6 10-6 8.77 6. 06 "13*' 1 239  MULTIPLE CURVILINEAR COVARIANCE ANALYSES GROUP  B,  DOUGLAS-FIR (NORMAL WOOD) COMPRESSION WOOD TOTAL DIFFERENCE FOR TESTING SLOPES  2  E  WHERE Y = OVENDRY WEIGHT OF TRACHEID IN  G,  X  X = TRACHEID LENGTH IN MM  SUMS DIFFERENCE FOR TESTING LEVELS COMBINED REGRESSION ** SIGNIFICANT AT THE 11 LEVEL  - GREEN CONDITION  least-squares fitted line for Douglas-fir  least-squares fitted line for Douglas-fir Compression Wood  —I— 1.96  3.92  4.90  Tracheid Length, mm  -| 5.88  Figure  14:  Tracheid tracheid secutive (NH4NO3) pression zation. tracheid  w e i g h t , i n c r e m e n t w i d t h a n d mean length p a t t e r n s across s i x conincrements of a f e r t i l i z e d D o u g l a s - f i r w h i c h p r o d u c e d comwood 4.years a f t e r f i r s t fertili(Each p o i n t r e p r e s e n t s t h e average weight or length f o r the increment).  Tracheid Weight, g- x l O  - 6  (Ovendry Condition)  Increment Width, mm-  Mean Tracheid Length, mm-  Figure  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 ) a n d 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 Douglas-fir.  135. HOLQ UREA  A.  Yr2==-0.24 02.3635;+S0.0 -997X2 0.103 10-6 2 0.948 1 0.008 N.S.  MULTIPLE CURVILINEAR COVARIANCE ANALYSES GROUP  DF  DOUGLAS-FIR  E  F  DF = 51  2 0 1  UREA FERTILIZED DOUGLAS-FIR  WHERE Y = OVENDRY WEIGHT OF TRACHEID  50  TOTAL  IN  2 5 1  DIFFERENCES FOR TESTING SLOPES  N.S.  SIMS  2 5 3  DIFFERENCES FOR TESTING LEVELS COMBINED REGRESSION  e.  x  X = TRACHEID LENGTH IN MM - GREEN CONDITION  2 5 4  N.S. NOT SIGNIFICANT AT THE 5% LEVEL  ion  <r least-squares fitted line for Douglos-fir  •o s c 0 oo-  ^least-squares fitted line for Douglas-fir, Urea fertilized  >.  •o O  0 0  g  •v  >  —I 1.96  1 2.94  I 5.88  —I  1 3.92  4.90  1 6.86  1.82  9.8c  X  Weic  ALPHA UREA •5  <u  f  0 r—  B,  to  GROUP  DF  DOUGLAS-FIR UREA FERTILIZED DOUGLAS-FIR TOTAL  DIFFERENCES FOR TESTING SLOPES SUMS DIFFERENCES FOR TESTING LEVELS COMBINED REGRESSION  F  13 99 6 232 50.721 23172,517 238  LEVEL N . S .  NOT SIGNIFICANT AT THE 5Z  Y R= 0.6502 0 .641 0X+0.0 .5 2264X .2== 0 . 9 4 2 ; S = 0 2 39 10"6 2  MULTIPLE CURVILINEAR COVARIANCE ANALYSIS  E  DF  WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G, X  N.S,  X - TRACHEID LEN3TH IN Ml  N.S,  - GREEN CONDITION  least-squares fitted line for Douglas-fir  ~leost-squares fitted line for Douglas-fir.Urea fertilized 2.94  3.92  T  4.90  Tracheid Length,mm-  ,A  Figure  16:  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) 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 ) a n d l e n g t h i n NPK treated and n o r m a l D o u g l a s - f i r .  HOLO NPK  Yr==-0.00 1 70 7.-+S 0 .069 W . 7 1 = 0 . 063 = 50 10-6 2  2  MULTIPLE CURVILINEAR COVARIANCE ANALYSIS  21 09 1 252 0 0.106.NS., 2512 0.581 253  GROUP  DF  DOUGLAS-FIR NPK FERTILIZED DOUGLAS-FIR TOTAL DIFFERENCES FOR TESTING SLOPES SUMS DIFFERENCES FOR TESTING LEVELS  F  N.S.  COMBINED REGRESSION  E  DF  WHERE Y = OVENDRY WEIGHT OF TRACHEID IN G. X  X = TRACHEID LENGTH IN MM - GREEN CONDITION  N,S. NOT SIGNIFICANT AT THE 5? LEVEL  ALPHA NPK  Y<==Q033 5 +S0 . C 0 , 8 6 1 ; 10 = 0.030 2  MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP  DF  DOUGLAS-FIR NPK FERTILIZED DOUGLAS-FIR  235  SUMS  IN G. X IO"  2  COMBINED REGRESSION  1 238  SIGNIFICANT AT THE V. LEVEL  6  X = TRACHEID LENGTH IN MM  237  DIFFERENCES FOR TESTING LEVELS "  WHERE Y = OVENDRY WEIGHT OF TRACHEID  39  TOTAL DIFFERENCES FOR TESTING SLOPES  EE  DF =  196  1.753  - GREEN CONDITION N.S.  N.S. NOT SIGNIFICANT AT THE 5% LEVEL  Tracheid Length, mm-  »A  Figure  17:  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 (NH4)2S04 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 NS04  Y'==-0.00 7, 56 887+S0.= 0676X 0.079 = 67 G, x 10-6 2  2  E  MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP  (NHi|)2S0i(  TOTAL FERTILIZED DOUGLAS FIR DIFFERENCES FOR TESTING SLOPES SUMS  DIFFERENCES FOR TESTING LEVELS COMBINED REGRESSION  N.S.  F  DF  DOUGLAS-FIR  26 06 1 262 70.309 N.S 261916.1)5 N.S 270  DF  WHERE Y = OVENDRY WEIGHT OF. TRACHEID  IN  X = TRACHEID LENGTH IN MM GREEN CONDITION  NOT SIGNIFICANT AT THE 57. LEVEL  ALPHA NS04  GROUP  DF  DOUGLAS-FIR  (NhtyjSQi)  F  196  FERTILIZED DOUSLAS-FIR TOTAL  DIFFERENCES FOR TESTING SLOPES SUMS DIFFERENCES FOR TESTING LEVELS COMBINED REGRESSION  12  IN G. X  13,631  210 1  DF  WHERE Y = OVENDRY WEIGHT OF TRACHEII  238 2  Y<==-0, 2 51 55 3,-+SEE 0.2599 X0. '019 0 . 8 = 13 10-6 2  MULTIPLE CURVILINEAR COVARIANCE ANALYSIS  21,103*  211  ' SIGNIFICANT AT THE 1% LEVEL  Tracheid Length,mm-  X " TRACHEID LENGTH IN MM GREEN CONDITION  .A  Figure  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 alpha-cellulose  (B)  skeletons)  treated  and  and l e n g t h  normal  i n NH4NO3  Douglas-fir.  HOLD HRNET  2-6  MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP  jf  DOUGLAS-FIR  201  Y = 0.6561 - 0.5187X + 0.1625 X2  p  R = 0.781; 2  FERTILIZED DOUGLAS FIR  NH4NO3  TOTAL  113  SUMS  COMBINED REGRESSION  DF = 149  2  56,192*'  1  x  IO  -6  X = TRACHEID LENGTH IN Ml - GREEN CONDITION  316  DIFFERENCES FOR TESTING LEVELS  E  IN G.  311  DIFFERENCES FOR TESTING SLOPES  SE = 0.113;  WHERE Y = OVENDRY WEIGHT OF TRACHEID  11,581"  317  ** SIGNIFICANT AT THE 1% LEVEL  least-squares fitted line for Dauglas-fir  •least-squares fitted line for Douglas-fir, N H 4 N O 3 fertilized  -r  2-94  1.96  6.86  —1 8.82  flLPHR HRNEY 2-6 Y = -0.0507 + 0.085D<2  MULTIPLE CURVILINEAR COVARIANCE ANALYSIS GROUP  DF  DOUGLAS FIR  NH1IO3  FERTILIZED  2  SUMS  = 0,072; DF = 115  E E  x  10-6  X = TRACHEID LENGTH IN Ml  305 2  S  IN G,  DOUGLAS-FIR 109  DIFFERENCES FOR TESTING SLOPES  = 0.909;  WHERE Y = OVENDRY WEIGHT CF TRACHEID  196  TOTAL  - GREEN CONDITION  51,519"  307  DIFFERENCES FOR TESTING LEVELS COMBINED REGRESSION  T  F  1  2.395 N.S.  308  N.S. NOT SIGNIFICANT AT THE 57. LEVEL **  SIGNIFICANT AT THE 12 LEVEL  least-squares fitted line for Douglas-fir  leost-squores filled line for Douglas-fir, NH4NO3  3.92  4.90  Tracheid Length, mm-  fertilized  —I  6.86  A P P E N D I C E S  APPENDIX DESCRIPTION  Araucaria cunninghamii Sequoia serapervirens Pinus lambertiana Podocarpus dacrydioides Picea sitchensis Taxus b r e v i f o l i a Juniperus virginiana Cephalotaxus wilsoniana ^Pseudotsuga  2 2  OF GROWTH INCREMENTS  Age From Pith Yr.  Species  menziesii  Pseudotsuga  menziesii  Pseudotsuga  menziesii  Pseudotsuga  menziesii  Pseudotsuga  menziesii  Pseudotsuga  menziesii  I  —  Wood Zone  INCLUDED  Increment Width mm  IN THE  Latewood Width mm  STUDY Measured Tracheid Length Average  ct  nun  Range  8  H H H H H H H S  2.4 2.5 1.4 3.6 4.4 2.3 3.4 6.5  0.4 0.5 0.3 0.6 1.4 0.6 0.8 0.8  7.42 5.97 • 6.29 4.59 4.70 2.14 2.37 2.13  5.10-9.79 4.20-8.50 3.75-8.12 2.92-6.04 2.92-5.84 1.44-2.96 1.42-2.99 1.70-2.56  10 20 80 150 300 400 73 83 20 21 22 23 24 25 15 16 15 16 15 16  H H H H H H H S H H H H S S S S H H H H  2.0 5.8 3.4 1.7 1.0 0.7 4.9 5.0 5.9 7.7 10.0 8.5 8.5 8.6 6.5 5.5 10. 5 10.7 7.7 7.5  0.9 2.4 1.4 0.7 0.3 0.2 3.5 3.2 2.5 3.7 3.3 2.4 2.6 3.7 1.5 1.1 2.0 2.0 1.1 0.9  2.37 3.97 5.46 5.92 4.96 5.89 2.84 2.89 2.92 2.50 2.51 2.40 2.81 2.30 3.31 2.80 2.39 2.54 2.23 2.39  1.88-2.71 2.14-4.55 3.96-6.67 3.33-7.08 3.13-6.88 3.45-7.08 0.85-3.27 1.99-3.27 1.20-3.80 1.45-4.20 1.65-3.95 1.90-4.20 1.65-3.77 1.05-3.85 2.70-3.98 2.13-3.56 1.85-3.13 1.99-3.13 1.42-2.99 1.99-3.56  36 70  -  (Continued)  -  140.  APPENDIX  Average  based  I  o n a minimum  (Continued)  o f 50 t r a c h e i d s  Heartwood Sapwood  Butt, l o g o f a 500-year-old Compression  wood  - Breast  tree height  disk  o f a 120-year-old  tree  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 fertilized Urea  fertilized  a t ages  21 a n d 22  - 12 mm i n c r e m e n t core from b r e a s t height, 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 a g e s 13 a n d 15 ( 2 0 0 l b / a c r e o f N , P 0 p r K 0 e q u i valents ) 2  NPK  fertilized  (NH^) SO^ 2  treej (500 l b / a c r e )  2  - 12 mm i n c r e m e n t core from b r e a s t height, 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 a g e s 13 a n d 15 ( 2 0 0 l b / a c r e o f N ^ - O ^ o r K . O equivalents)  fertilized  - 12 mm i n c r e m e n t core from breast height, 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 ( 2 0 0 l b / a c r e o f N , P 0 o r K 0 equivalents) 2  5  2  APPENDIX I I SUMMARY OF H O L O C E L L U L O S E L I G N I N I N SOME OF THE SAMPLES  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 OF S T U D I E D (BASED ON OVENDRY, E X T R A C T I V E - F R E E WOOD)  Description of Species  Species  Increment No. F r o m Pith  Portion i n the Increment  EW LW EW LW LW EW LW EW ' LW LW  Pseudotsuga  menziesii  500-year-old  80  Pseudotsuga  manziesii  NH .NO,  23  fertilized  Pseudotsuga menziesii Sequoia sempervirens  C o m p r e s s i o n wood M a t u r e wood  73  Picea  sitchensis  Mature  wood  36  Pinus  lambertiana  Mature  wood  —  a  Each c y c l e followed  b  No. i n p a r e n t h e s i s  EW LW  Earlywood Latewood  -  No. o f Cycles  3  3 5 3 5 5 3 5 3 5 5  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 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 ) indicates  percentage of r e s i d u a l  lignin  Holocellulose Y i e l d %  7-4 . 8 ( 1 . 3 0 ) 74 .1 74 .3 73 .9 61 .2 70 .1 68 .9 74 .0 73 .4 72 . 2 ( 1 . 53)  acid,  Alpha-Cellulose Y i e l d %  b  45. 46. 44. 45. 38. 43. 44. 46. 47. 46.  2 ( 0 . 27) 3 9 8 4 3 2 0 . 5 ( 0 . 29) 4  142.  APPENDIX I I I M O I S T U R E CONTENT OF 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 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 C T H ROOM FOR OVER ONE MONTH  Earlywood/ Latewood  Species  Holocellulose Moisture Content  Alpha-Cellulose Moisture Content  Q.  O.  "5  Araucaria Sequoia Pinus  cunninghamii  sempervirens  lambertiana  Podocarpus  dacrydiodes  Picea  sitchensis  Taxus  brevifolia  Juniperus  virginiana  Cephalotaxus  wilsoniana  Pseudotsuga I n c r e m e n t 10 i n c r e m e n t 20  menziesii  Increment  80  I n c r e m e n t 150 Increment  3 00  I n c r e m e n t 400 2  Pseudotsuga  menziesii  Pseudotsuga I n c r e m e n t 20 I n c r e m e n t 21  menziesii  3  Increment  22  EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW  10.66 10.87 9.86 9.79 9.80 10.04 9.74 9.81 10.58 10.73 10.00 10.03 9.80 10.07 10.20 10.20 10.73 10.01 10.08 9.98 9.99 10.05 10.88 10.98 9.99 10.01 10.59 10.11 10.26 10.28 10.34 10.35 10.31 10.34 10.19 10.18  *6  10.15 10.38 9.52 9.62 9.83 10.12 9.47 9.52 9.8 0 9 . 54 9.65 9.84 9.60 9.96 9.31 9.31 9.75 10.10 10.11 9.71 10.07 9. 94 10.16 10.24 9.90 9.73 9.92 9.96 9.52 9.54 9.43 9.46 9.61 9.64 9.40 9.46  (Continued)  143.  APPENDIX I I I ( C o n t i n u e d )  Earlywood/ Latewood  Species  Holocellulose Moisture Content  Alpha-Cellulose Moisture Content a *o  Q,  *5  Increment  23  Increment  24  Increment  25  Pseudotsuga I n c r e m e n t 15 I n c r e m e n t 16  menziesii  Pseudotsuga I n c r e m e n t 15 I n c r e m e n t 16  menziesii  Pseudotsuga I n c r e m e n t 15 I n c r e m e n t 16  menziesii  l o g of a  5 0 0 -y e a r - o l d  1.  Butt  2.  C o m p r e s s i o n wood  3.  NH.NO_ 4 j  4.  Urea  10.18 10.28 10.20 10.18 10.19 10.88 9.85 9. 86 9.88 9.85 10.47 10.50 10.42 10.48 10.62 10.59 10.58 10.57  10.05 9.86 9.79 9.78 9.88 9.90 9.49 9.52 9.53 9.52 9.56 9.58 9.50 9.52 9.64 9.62 9.59 9.61  tree  - B r e a s t '.h e i g h t d i s k  fertilized -  fertilized  EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW EW LW  Breast height fertilized at  of a  d i s k of a g e s 21  120-year-old  tree  a 29-year-old tree; a n d 22 (500 l b / a c r e  - 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 a g e s 13 a n d 15 (200 l b / a c r e o f N,P 0,.orK 0 equivalents) 2  5.  NPK  fertilized  - 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 a g e s 13 a n d 15 ( 2 0 0 l b / a c r e o f - N , P 0 o r K 0 equivalents) 2  6.  (NH^) SO^ 2  fertilized  5  2  - . 1 2 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 0 o r K 0 equivalents) 2  2  

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