Professional Documents
Culture Documents
R. J. THOMAS
Department of Wood and Paper Science, North Carolina State University,
Raleigh, N.C. 27607
Figure 1. End view of a log showing both the wood and bark.
Note the small total volume occupied by the bark. The central,
dark portion of the wood is heartwood, and the outer ring of
light colored wood is sapwood.
f o o d - s t o r a g e c e l l s , t h u s r e c o g n i t i o n o f h e a r t w o o d based on c o l o r
i s not always p o s s i b l e . I t s h o u l d be a p p a r e n t t h a t t h e v a s t ma-
j o r i t y o f c e l l s w h i c h c o n s t i t u t e t h e wood p o r t i o n o f a l i v i n g
t r e e a r e dead* The sapwood zone c o n t a i n s t h e o n l y l i v i n g c e l l s
f o u n d i n mature wood and t h e y c o n s t i t u t e , d e p e n d i n g upon s p e c i e s ,
f r o m 10 t o kO p e r c e n t o f t h e sapwood volume. As h e a r t w o o d i s
formed t h e s e l i v i n g , f o o d - s t o r a g e c e l l s d i e , t h u s t h e h e a r t w o o d
c o n t a i n s o n l y dead c e l l s .
T r e e s a r e c l a s s i f i e d i n t o two m a j o r g r o u p s termed s o f t w o o d s
(gymnosperms) and hardwoods ( a n g i o s p e r m s ) . The b o t a n i c a l b a s i s
f o r c l a s s i f i c a t i o n i s w h e t h e r o r n o t t h e t r e e seed i s naked as i n
s o f t w o o d s o r c o v e r e d a s i n hardwoods. A more f a m i l i a r c l a s s i f i -
c a t i o n , w h i c h w i t h some e x c e p t i o n s i s v a l i d , i s based on t h e r e -
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t e n t i o n o f l e a v e s by s o f t w o o d s o r t h e s h e d d i n g o f l e a v e s by h a r d -
Publication Date: June 1, 1977 | doi: 10.1021/bk-1977-0043.ch001
woods. Thus t h e s o f t w o o d s a r e o f t e n r e f e r r e d t o as e v e r g r e e n
t r e e s and hardwood as d e c i d u o u s t r e e s . The m a j o r d i f f e r e n c e w i t h
r e g a r d t o wood anatomy i s t h e p r e s e n c e o f v e s s e l s i n hardwoods.
V e s s e l s a r e s t r u c t u r e s composed o f c e l l s c r e a t e d e x c l u s i v e l y f o r
t h e c o n d u c t i o n o f w a t e r . S o f t w o o d s l a c k v e s s e l s b u t have c e l l s
termed l o n g i t u d i n a l t r a c h e i d s w h i c h p e r f o r m a d u a l r o l e o f c o n -
d u c t i o n and s u p p o r t .
The terms s o f t w o o d and hardwood a r e n o t t o be t a k e n a s a
measure o f h a r d n e s s s i n c e some hardwoods a r e s o f t e r t h a n many
softwoods. F o r t h e c o m m e r c i a l l y i m p o r t a n t d o m e s t i c woods, t h e
a v e r a g e s p e c i f i c g r a v i t y f o r s o f t w o o d s i s .41 w i t h a range o f .29
t o .60. Hardwoods a v e r a g e .50 and v a r y f r o m .32 t o .81.
Growth r i n g s , o r a n n u a l i n c r e m e n t s , as seen on t h e end o f a
s o f t w o o d l o g a r e d e p i c t e d i n F i g u r e 2. Growth r i n g s a r e d e t e c t a -
b l e due t o d i f f e r e n c e s i n t h e wood p r o d u c e d e a r l y and l a t e i n t h e
g r o w i n g s e a s o n . The wood p r o d u c e d e a r l y , c a l l e d e a r l y w o o d o r
s p r i n g w o o d , i s c o n s i d e r a b l y l i g h t e r i n c o l o r t h a n t h e wood termed
l a t e w o o d o r summerwood w h i c h i s p r o d u c e d l a t e i n t h e g r o w i n g
s e a s o n . The c o l o r d i f f e r e n c e i s due p r i m a r i l y t o t h e d i f f e r e n t
k i n d s o f c e l l s produced e i t h e r e a r l y o r l a t e i n t h e growing sea-
son. F i g u r e 3 d e p i c t s t h r e e g r o w t h r i n g s and p a r t o f a f o u r t h .
Note t h a t a t t h i s low m a g n i f i c a t i o n y o u c a n d e t e c t t h e i n d i v i d u a l
c e l l s w h i c h c o n s t i t u t e t h e s p r i n g w o o d a r e a ; h o w e v e r , t h e summer-
wood zone a p p e a r s m e r e l y as a d a r k z o n e . A t a h i g h e r m a g n i f i c a -
t i o n , a s shown i n F i g u r e 4, i n d i v i d u a l c e l l s o f b o t h s p r i n g w o o d
and summerwood a r e e v i d e n t . The s p r i n g w o o d c e l l s have a l a r g e
c r o s s - s e c t i o n a l a r e a w i t h t h i n w a l l s and a l a r g e , open c e n t e r .
The l a r g e , open c e n t e r , c a l l e d t h e lumen, p e r m i t s e f f i c i e n t c o n -
d u c t i o n o f w a t e r . The summerwood c e l l s have a s m a l l e r c r o s s -
s e c t i o n a l a r e a , t h i c k e r w a l l s and a s m a l l e r lumen t h a n s p r i n g w o o d
cells. O b v i o u s l y , t h i s type o f c e l l provides s u b s t a n t i a l support
f o r t h e t r e e stem b u t i s n o t as e f f i c i e n t as s p r i n g w o o d c e l l s i n
conduction. In some s o f t w o o d s p e c i e s t h e g r a d a t i o n between
s p r i n g w o o d and summerwood i s g r a d u a l and f e w e r summerwood c e l l s
a r e p r o d u c e d . When t h i s o c c u r s , i t i s o f t e n d i f f i c u l t t o d i s t i n -
g u i s h g r o w t h i n c r e m e n t s a s t h i s t y p e o f wood p o s s e s s e s a more
uniform s t r u c t u r e .
As i n d i c a t e d e a r l i e r , hardwoods a r e c h a r a c t e r i z e d by t h e
presence o f vessels o r pores. Vessels a r e c e l l s which occupy a
l a r g e c r o s s - s e c t i o n a l a r e a and c a n i n most s p e c i e s be d e t e c t e d
w i t h t h e u n a i d e d e y e . The wood o f hardwood t r e e s i s c l a s s i f i e d
as eî t h e r r i n g - p o r o u s o r d i f f u s e - p o r o u s depend i ng upon t h e a r -
rangement o f t h e v e s s e l s. I η a t y p i c a l r i n g - p o r o u s wood, as i l
l u s t r a t e d i n F i g u r e 5, t h e v e s s e l s formed i n t h e s p r i ngwood a r e
c o n s i d e r a b l y l a r g e r t h a n t h o s e formed i n t h e summerwood. F i g u r e
6 r e v e a l s a d î f f u s e - p o r o u s wood i n w h i c h t h e v e s s e l s a r e e s s e n -
t i a l l y t h e same s i z e t h r o u g h o u t t h e g r o w t h r i n g . As a r e s u l t , i n
many d i f f u s e - p o r o u s woods i t i s d i f f i c u l t t o d i s t i n g u i s h g r o w t h
rings. A c o m p a r i s o n o f F i g u r e s 3» 5 and 6 w i l l r e v e a l t h e m a j o r
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o f v e s s e l s and t h e r e l a t i v e l y un i f o r m a p p e a r a n c e o f t h e s o f t w o o d
i n F i g u r e 3, compared t o t h e hardwoods d e p i c t e d i n F i g u r e s 5 and
6.
Wood r a y s a r e f o u n d i n a l 1 s p e c i e s o f wood and cons i s t o f
ribbon-1ike aggregations o f food-storing c e l l s extending i n the
t r a n s v e r s e d i r e c t i o n from the bark toward t h e c e n t e r o f t h e t r e e .
In t h e c r o s s - s e c t i o n a l v i e w , r a y s t a k e t h e f o r m o f 1 i n e s o f v a r y -
i n g w i d t h r u n n i n g a t r i g h t a n g l e s t o t h e g r o w t h r i n g s ( F i g u r e s 3»
4, 5 and 6 ) .
S o f t w o o d Anatomy
The anatomy o f s o f t w o o d s w i l l be d e s c r i b e d f i r s t b e c a u s e i t
i s l e s s c o m p l e x t h a n hardwoods. The two mai η e e l 1 t y p e s wh i c h
c o n s t i t u t e s o f t w o o d s a r e t r a c h e i d s , w h i e h c o n d u c t and s u p p o r t ,
and parenchyma w h i c h s t o r e f o o d . T h e s e two e e l 1 t y p e s c a n be
f u r t h e r c l a s s i f i e d as t o t h e i r o r i e n t a t i o n , t h a t i s l o n g i tud i n a l
or transverse. C e l l s o r i e n t e d i n t h e l o n g i tud înal d i r e c t i o n have
t h e l o n g a x i s o f t h e e e l 1 o r i e n t e d p a r a 1 l e i t o t h e v e r t îcal a x i s
o f t h e t r e e t r u n k w h e r e a s t r a n s v e r s e l y o r i e n t e d e e l 1 s have t h e i r
long a x i s a t r i g h t a n g l e s t o t h e v e r t i c a l a x i s o f t h e t r e e stem.
A g r e a t l y s împli f i e d model o f s o f t w o o d anatomy c a n be made by
g l u i ng t o g e t h e r a g r o u p o f s o d a s t r a w s a l o n g t h e i r l e n g t h and
d î s p e r s i ng t h r o u g h o u t t h i s g r o u p m a t c h e s , l a i d end t o end w i t h
t h e l o n g a x i s o f t h e matches o r i e n t e d a t r i g h t a n g l e s t o t h e l o n g
axi s o f t h e soda s t r a w s . The s o d a s t r a w s r e p r e s e n t t h e l o n g i t u -
d i n a l l y o r i e n t e d t r a c h e i d s w h i c h o c c u p y a b o u t 30% o f t h e vo1ume
w h e r e a s t h e matches r e p r e s e n t t r a n s v e r s e l y o r i e n t e d parenchyma
e e l Is o c c u p y i n g a b o u t 10% o f t h e vo1ume. The t r a n s v e r s e l y o r i -
e n t e d parenchyma a r e t h e e e l 1 s w h i c h c o n s t i t u t e wood r a y s . As
t h e matches i n t h e model a r e cons i d e r a b l y s h o r t e r t h a n t h e soda
s t r a w s , s o parenchyma e e l 1 s a r e c o n s i d e r a b l y s h o r t e r t h a n l o n g i -
tudinal tracheids.
S i n c e l o n g i t u d i n a l t r a c h e i d s c o n s t i t u t e a b o u t 30% o f t h e
vo1ume and a r e t h e r e f o r e l a r g e l y r e s p o n s i b l e f o r t h e r e s u l t i n g
phys i c a l and chemi c a l p r o p e r t i e s o f s o f t w o o d s , a d e t a i l e d
d e s c r i p t i o n o f t h e i r anatomy i s m e r i t e d . F i g u r e 3» s h o w i n g a
smal1 b l o c k o f p i ne magni f i e d 50 t i m e s , c l e a r l y î11ustrates t h a t
s p r i n g w o o d and summerwood 1 o n g i t u d i n a l t r a c h e i d s make up t h e
l a r g e s t vo1ume o f t h e wood. Note t h a t i n add i t i o n t o t h e c r o s s -
s e c t i o n a l v i e w , two l o n g i t u d i n a l v i e w s a r e e x p o s e d . Transversely
o r i e n t e d parenchyma e e l I s , w h i c h make up t h e r a y s , c a n a l s o be
s e e n i n t h e t r a n s v e r s e and l o n g i t u d i n a l v i e w s . I η the cross-
s e c t i o n a l v i e w , t h r e e c o m p l e t e and two p a r t i a l g r o w t h r i n g s a r e
depicted.
The l o n g i t u d i n a l t r a c h e i d i s a b o u t 100 t i m e s l o n g e r t h a n
w i d e . D e p e n d i n g on s p e c i e s , most d o m e s t i c s o f t w o o d s have l o n g i
t u d i n a l t r a c h e i d s r a n g i n g f r o m 3 t o 5 mm i n l e n g t h . Redwood has
t h e l o n g e s t t r a c h e i d s , up t o 7.3 mm, and c e d a r t h e s h o r t e s t ,
a b o u t 1.18 mm i n l e n g t h . The w i d t h o f l o n g i t u d i n a l t r a c h e i d s f o r
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Publication Date: June 1, 1977 | doi: 10.1021/bk-1977-0043.ch001
d o m e s t i c s p e c i e s r a n g e s f r o m 20 ym f o r c e d a r up t o 80 ym f o r
redwood.
S i n c e t h e l o n g i tud i n a l t r a c h e i d i s a l o n g , t h i n, c y 1 i n d r i c a l ,
t u b e - 1 i ke e e l 1, i t s a p p e a r a n c e depends upon how i t i s ν i ewed.
For example, i f t h e c e l l i s c u t a t r i g h t a n g l e s t o i t s long a x i s ,
t h e c r o s s - s e c t i o n a l v i e w i s e x p o s e d on t h e c u t s u r f a c e . The
c r o s s - s e c t i o n a l v i e w e x p o s e d i n F i g u r e 3 r e v e a l s î η t h e sprî ng-
wood a s q u a r e o r p o l y g o n a l s h a p e predomi natîng wh i l e i n t h e sum-
merwood a more r e c t a n g u l a r shape i s a p p a r e n t . I η the longi tudi
nal , r a d i a l p l a n e v i e w , t h e ends o f sprî ngwood t r a c h e i d s a r e
rounded, w h i l e i n t a n g e n t i a l view they a r e p o i n t e d ( F i g u r e 7 ) .
The ends o f summerwood t r a c h e i d s a r e p o i n t e d i n b o t h r a d i a l and
t a n g e n t i a l views (Figure 7 ) .
11 s h o u l d be o b v i o u s t h a t t h e s t r u c t u r e o f t h e l o n g i t u d i n a l
t r a c h e i d i s w e l l s u i ted t o perform t h e dual r o l e s o f conduct ion
and s u p p o r t . S i nee w a t e r i s t r a n s l o c a t e d up t h e t r e e v i a t h e
t r a c h e i d s , the o r i e n t a t ion o f the long a x i s o f the t r a c h e i d par-
a i l e i t o t h e v e r t i c a l s t e m p e r m i t s a l o n g e r passageway p r i o r t o
i n t e r r u p t i o n by a c e l l w a l l . The r i g i d c e l l w a l l s , o f v a r y i n g
t h i c k n e s s , p r o v i d e adequate support.
Of t h e s e v e r a l t y p e s o f m a r k i ngs f o u n d on t h e l o n g i t u d i n a l
tracheîd w a l l ( F i g u r e s 7 and 8 ) , t h e most o b v i o u s a r e t h e c i r c u -
l a r dome-1i ke s t r u c t u r e s c a l l e d b o r d e r e d p i t s . Also depicted are
c l u s t e r s o f egg shaped p i t s whi ch î n t e r c o n n e c t l o n g i t u d i n a l t r a -
c h e i d s t o t r a n s v e r s e l y o r i e n t e d parenchyma c e l l s . The s t r u c t u r e
o f t h e b o r d e r e d p i t f a c i l i t a t e s l i q u i d f l o w between c e l l s . An
o p e n i ng termed t h e p i t a p e r t u r e i s l o c a t e d i n t h e c e n t e r o f t h e
dome-1ike s t r u c t u r e w h i c h i s c a l l e d t h e p i t b o r d e r ( F i g u r e 9 ) .
Removal o f t h e b o r d e r r e v e a l s t h e p i t membrane w h i c h r e s e m b l e s a
w h e e l w i t h a hub and s p o k e s ( F i g u r e 1 0 ) . The p o r t i o n o f t h e mem-
brane s i m i l a r t o t h e p a r t o f t h e wheel w i t h spokes i s c a l l e d t h e
margo. N o t e t h e many o p e n i n g s i n t h e margo t h r o u g h w h i c h l i q u i d
can f l o w . T h e c e n t r a l p o r t i o n , r e s e m b l i ng t h e hub o f a w h e e l ,
has no d e t e c t a b l e open i n g s and i s termed t h e t o r u s . S i nee e a c h
b o r d e r e d p i t wi t h i η a c e l l u s u a l l y has a c o m p l i m e n t a r y p i t i n t h e
c o n t i g u o u s e e l 1, 1 i q u i d c a n f l o w f r o m t h e lumen t h r o u g h t h e p i t
Wood Science
b o r d e r and t h e e x t r e m e l y t i g h t s e a l w h i c h e x i s t s between t h e t o r -
us and p i t b o r d e r . A s p i r a t i o n o f t h e p i t membrane o c c u r s a s w a t -
e r i s removed f r o m t h e c e l l . In t h e l i v i n g t r e e , a s p i r a t i o n p r e -
v e n t s , i n t h e e v e n t a t r e e i s wounded, a i r e m b o l i s m s f r o m o c c u r -
r i n g t h r o u g h o u t t h e t r e e and i n t e r r u p t i n g a l 1 w a t e r c o n d u c t i o n .
P i t membrane a s p i r a t i o n , w h i c h o c c u r s d u r i ng t h e d r y i ng o f wood,
r e d u c e s wood permeab î1i t y as 1 i q u i d f l o w between e e l 1 s i s p r e -
vented. Thus any p r o c e s s i ng o f wood î n v o l v i ng t h e pénétrât i o n o f
l i q u i d s a f t e r d r y i n g i s more d i f f i c u l t .
The o t h e r m a j o r e e l 1 w a l l s t r u c t u r e found on l o n g i t u d i n a l
t r a c h e i d s i s termed a r a y c r o s s i ng and i s î11ustrated i n F i g u r e s
7 and 8. Ray c r o s s î ngs c o n s i s t o f p i t s w h i c h î n t e r c o n n e c t l o n g i -
tud î n a l t r a c h e i d s t o r a y parenchyma. Due t o t h e d i v e r s e s t r u c -
t u r e o f r a y c r o s s i n g p i t s they a r e extremely u s e f u l i n t h e iden-
t i f i c a t i o n o f wood and wood f i b e r s . However, s i nee i d e n t i f i c a -
t i o n i s beyond t h e s c o p e o f t h i s r e v i e w , a descrî pt i o n o f t h e
d i f f e r e n t t y p e s o f p i t s found i n r a y c r o s s i n g s i s n o t i n c l u d e d .
F i g u r e s 8 and 15 r e v e a l r a y c r o s s i ng p i t s as s e e n f r o m t h e
i n s i d e o f l o n g i t u d i n a l t r a c h e i d s . The c o n s i d e r a b l y h i g h e r magni-
f i c a t i o n i n F i g u r e 15 shows a s o l i d p i t membrane. O p e n i n g s i n
t h e p i t membrane w o u l d e x p o s e t h e c y t o p l a s m t o t h e h o s t île e n v i -
ronment o f t h e l o n g i t u d i n a l t r a c h e i d lumen and r e s u l t i n t h e
d e a t h o f t h e parenchyma e e l 1. T h u s , t h e membranes a r e s o l i d and
do n o t p r o v i d e a passageway f o r f r e e l i q u i d f l o w .
Hardwood Anatomy
T h u s , most hardwood s p e c i e s c o n t a i n f o u r t y p e s o f c e l l s , v e s s e l
s e g m e n t s , f i b e r s , t r a n s v e r s e and a x i a l parenchyma, w h e r e a s most
s o f t w o o d s p e c i e s p o s s e s s two t y p e s ; l o n g i t u d i n a l t r a c h e i d s and
t r a n s v e r s e parenchyma.
V e s s e l s a r e s t r u c t u r e s u n i q u e l y des i g n e d t o c a r r y o u t t h e
conduct ion r o l e . A v e s s e l cons i s t s o f i n d i v i d u a l v e s s e l e e l 1 s
s t a c k e d one on t o p o f t h e o t h e r i n t h e v e r t i c a l d i r e c t i o n o f t h e
s t e m ( F i g u r e 6 ) . V e s s e l l e n g t h s up t o t h r e e m e t e r s have been
n o t e d . The v e s s e l c e l l s o r s e g m e n t s , w h i c h c o n s t i t u t e a v e s s e l ,
d i f f e r w i d e l y i n t h e i r s i z e and s h a p e . The l e n g t h o f v e s s e l s e g -
ments v a r i e s f r o m 0.18 mm t o 1.3 mm and f r o m 20 ym t o 330 ym i n
width. The s h o r t e s t and w i d e s t v e s s e l segments a r e f o u n d i n t h e
s p r î ngwood o f r i n g - p o r o u s woods where o f t e n t h e w i d t h o f t h e v e s -
s e l segment i s g r e a t e r t h a n t h e l e n g t h . However, i n d i f f u s e -
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Publication Date: June 1, 1977 | doi: 10.1021/bk-1977-0043.ch001
p o r o u s woods t h e v e s s e l segments a r e u s u a l l y 8 t o 10 t i m e s l o n g e r
than they a r e wide. Note i n F i g u r e s 5 and 6 t h a t v e s s e l segments
i n c r o s s - s e c t i o n a l v i e w have a more o r l e s s rounded shape. Iη
t h i s v i e w t h e y a r e o f t e n ca1 l e d p o r e s whi ch i s a t e r m o f c o n v e n
i e n c e t o d e s c r i b e t h e c r o s s s e c t i o n o f a v e s s e l . As s e e n l o n g i
t u d i n a l l y i n F i g u r e 16, t h e y range f r o m drum-shaped t o b a r r e l -
shaped t o o b l o n g , 1î n e a r - s h a p e d e e l 1 s.
Two o b v i o u s s t r u c t u r a 1 f e a t u r e s o f v e s s e l segments a r e p e r -
forât i o n p l a t e s and p i t s . P e r f o r a t i o n p l a t e s a r e d i stî net open-
i n g s f o u n d a t b o t h ends o f t h e v e s s e l segment w h i c h l e a d t o a d j a -
cent v e s s e l segments. In some s p e c i e s a number o f o p e n i n g s a r -
ranged p a r a l l e l t o e a c h o t h e r f o r m t h e p e r f o r a t i o n p l a t e . These
a r e termed s c a l a r i f o r m perforât i o n p l a t e s . A s i n g l e o p e n i ng i s
c a l led a s impie p e r f o r a t i o n p l a t e . B o t h t y p e s o f perforât i o n
p l a t e s a r e î11ustrated i n F i g u r e 16. The c r e a t i o n o f o p e n i ngs
between v e s s e l segments p r o v i d e s an e l o n g a t e d t u b e l i k e s t r u c t u r e
of considerable length h i g h l y s u i t e d f o r the l o n g i t u d i n a l trans-
l o c a t i o n o f water.
When v e s s e l s e n d , t h e y r a r e l y do s o i n i s o l a t i o n b u t r a t h e r
among a g r o u p o f v e s s e l s. T r a n s l o c a t i o n c o n t i nues i n t o t h e a d j a -
cent v e s s e l s v i a the î n t e r v e s s e l pi t s . These pi t s d i f f e r from
s o f t w o o d b o r d e r e d p i t s i n t h a t t h e y l a c k a t o r u s and o p e n i n g s
l a r g e enough t o be r e a d i l y d e t e c t e d w i t h an e l e c t r o n m i c r o s c o p e .
F i g u r e 17 d e p i c t s a t y p i ca1 i n t e r v e s s e l p i t membrane. D i f f e r e n t
a r r a n g e m e n t s o f i n t e r v e s s e l p i t s c a n be d e t e c t e d and a r e u s e f u l
i n t h e i d e n t i f i c a t i o n o f hardwood s p e c i e s .
The l o n g i tud i n a l e e l 1 t y p e s r e s p o n s i b l e f o r t h e s u p p o r t r o l e
i n hardwoods a r e f i b e r s . F i b e r s a r e t h i c k - w a l l e d , e l o n g a t e d e e l Is
w i t h c l o s e d pointed ends. I t s h o u l d be n o t e d t h a t f r e q u e n t l y t h e
1 1
term " f i b e r s i s used l o o s e l y f o r a l 1 t y p e s o f wood e e l 1 s.
However, s p e c i f i c a l l y i t r e f e r s o n l y t o t h o s e e e l 1 t y p e s f o u n d i n
hardwoods w h i c h meet t h e above d e f i n i t i o n . F i b e r s range i n l e n g t h
f r o m 0.7 mm t o 3 mm w i t h an a v e r a g e s i i g h t l y l e s s t h a n 2 mm f o r
domestic species. i n d i a m e t e r , an a v e r a g e o f l e s s t h a n 20 ym can
be e x p e c t e d . The p e r c e n t a g e o f t h e volume o f wood o c c u p i e d by
f i b e r s v a r i e s cons i d e r a b l y . I η sweetgum, f i b e r s c o n s t i t u t e o n l y
t r a n s v e r s e p a r e n c h y m a , r a n g e f r o m one t o thî r t y - p l u s e e l 1 s w i d e .
The r a y i 1 1 u s t r a t e d i n F i g u r e 18 i s s e v e n e e l 1 s w i d e . Thus t h e
h i g h e r parenchyma volume i s due t o w i d e r r a y s and t h e a d d i t i o n a l
p r e s e n c e o f a x i a l parenchyma w h i c h i s r a t h e r r a r e i n s o f t w o o d
species.
Based on t h e wood a n a t o m i c a l d e s c r i p t i o n s p r e s e n t e d , i t i s
o b v i o u s t h a t hardwoods and s o f t w o o d s d i f f e r cons i d e r a b l y f r o m
e a c h o t h e r . F o r e x a m p l e , v e s s e l s a r e p r e s e n t i n hardwoods and
absent i n softwoods. I η hardwoods more e e l 1 t y p e s , s h o r t e r e e l 1s,
more parenchyma and a more v a r i a b l e a r r a n g e m e n t o f e e l 1 t y p e s
o c c u r . The r e l a t i ve u n i f o r m i t y o f s o f t w o o d anatomy i s t h e r e s u l t
of t h e preponderance o f a s i n g l e type c e l l , the l o n g i t u d i n a l
tracheid.
C e l l Wall Structure
A l t h o u g h some v a r i a b i 1 i t y e x i s t s , t h e i n t e r n a l e e l 1 w a l l
s t r u c t u r e d e s c r i b e d below r e p r e s e n t s t h e t y p i c a l s t r u c t u r e f o u n d
i n most woody p l a n t c e l l s . A t t h e t i m e t h e c e l l i s p r o d u c e d by
c e l l d i v i s i o n , i t cons i s t s o f a p r i m a r y w a l l wh i c h i s c a p a b l e o f
e n l a r g i ng b o t h l o n g i t u d i n a l l y and t r a n s v e r s e l y . A f t e r the eel 1
r e a c h e s f u l 1 s i z e , a s e c o n d a r y w a l l i s depos î t e d î n t e r n a l t o t h e
p r i m a r y w a l l add i ng t h i c k n e s s and r i g i d i t y t o t h e e e l 1 w a l l .
F i g u r e 19 i 1 1 u s t r a t e s t h e e e l 1 w a l I s o f two m a t u r e , c o n t i g u o u s
eel 1 s from a softwood spec i e s . Note t h e t h r e e d i s t i n e t 1 a y e r s
w h i c h make up t h e c e l l w a l l . A d j a c e n t t o and on e a c h s i d e o f t h e
middle l a m e l l a , a primary w a l l from both c e l l s i s p r e s e n t .
However, t h i s w a l 1 i s t o o t h î η t o be eas i l y o b s e r v e d . T h e r e f o r e ,
the t e r m compound m i d d l e 1amel1 a, w h i c h r e f e r s t o t h e m i d d l e l a
m e l l a and t h e two p r i mary wa11 s, î s o f t e n ut i 1 i z e d . A d j a c e n t t o
the compound m i d d l e l a m e l 1 a and e a s i l y d e t e c t e d i s t h e f i r s t 1ay-
e r o f t h e s e c o n d a r y w a l 1 termed t h e S^ l a y e r . The n e x t and c e n -
t r a l l a y e r , w h i c h i s t h e l a r g e s t , i s c a l l e d S^. The înnermost
l a y e r a d j a c e n t t o t h e lumen i s termed
The t o t a l e e l 1 wa11 t h i c k n e s s i s l a r g e l y c o n t r o l l e d by t h e
t h i c k n e s s o f t h e $ l a y e r . T h a t i s, t h i c k - w a l l e d e e l 1 s r e s u l t
9
Table I
T h i c k n e s s o f V a r i o u s C e l l Wal1 L a y e r s
and M i c r o f i b r i 1 A n g l e Wi t h î η t h e L a y e r s
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Publication Date: June 1, 1977 | doi: 10.1021/bk-1977-0043.ch001
Relative Average
Thickness Angle o f
Wal1 Layer (%) MicrofibriIs
D e t e c t i o n o f t h e v a r i o u s e e l 1 w a l 1 l a y e r s i s due t o t h e
p r e s e n c e o f m i c r o f i b r i 1 s wh i c h a r e o r i e n t e d a t d i f f e r e n t a n g l e s
wi t h i η e a c h l a y e r . A m i c r o f i b r i 1 i s t h e bas i c n a t u r a l l y o c c u r -
r i ng un i t wh î c h c a n be eas i l y s e e n wi t h an e l e c t r o n m i c r o s c o p e .
Note i n F i g u r e 21 t h e s t r i n g - l i k e a p p e a r a n c e o f t h e m i c r o f i b r i I s .
Q
In s i z e , m i c r o f i b r i I s range f r o m 1 0 0 t o 3 0 0 A i n d i a m e t e r , whereas
t h e i r l e n g t h has n o t been d e t e r m i n e d . Wi t h i η e a c h o f t h e e e l 1
wal1 l a y e r s , the m i c r o f i b r i 1 s a r e o r i e n t e d at di f f e r e n t angles t o
t h e l o n g a x i s o f t h e e e l 1. T a b l e I i n d i c a t e s t h e a v e r a g e m i c r o -
f i b r i 1 o r i e n t a t i o n wi t h i η t h e v a r i o u s e e l 1 w a l 1 l a y e r s . Figure
22 p r e s e n t s an i d e a l i z e d d r a w i n g o f t h e microfibrî1 o r i e n t a t i o n
wi t h i η t h e v a r i o u s e e l 1 w a l 1 l a y e r s .
Figure 20. Cell walls of a vessel (V) and adjacent fiber (F) in
cross-sectional view. Note the very large S layer in the thick-
B
13β00χ
Table ! I
Softwoods Hardwoods
Cel1ulose 42 ± 2 45 ± 2
Hemicel1ulose 27 ± 2 30 ± 5
Lignî η 28 ± 3 20 ± 4
Extractî ves 3 ± 2 5 ± 3
C e l 1 u l o s e i s a 1 i n e a r polymer o f anhydro-D-glucopyranose
uni t s 1i nked by $-(l->4) g l y c o s id i c bonds. The number o f g l u c o s e
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certaî n.
H e m i c e l 1 u l o s e s , 1i ke c e l 1 u l o s e , a r e p o l y m e r s o f a n h y d r o s u g a r
uni t s . They d i f f e r f r o m c e l 1 u l o s e î n t h a t a g i ven h e m i c e l 1 u l o s e
m o l e c u l e may c o n t a i n s e v e r a l d i f f e r e n t s u g a r un i t s . In add i t i o n ,
h e m i c e l 1 u l o s e s a r e u s u a l 1 y b r a n c h e d m o l e c u l e s containîng o n l y 150
t o 200 s u g a r u n i t s . H e m i c e l l u l o s e s found î n wood a r e p o l y m e r s o f
D - g l u c o s e , D - g a l a c t o s e , D - x y l o s e , D-mannose, L - a r a b i nose and 4-0-
methy1-D-glucuronic a c i d . The h e m i c e l 1 u l o s e s a l o n g wi t h 1 i g n i n
s u r r o u n d t h e c r y s t a l 1i ne c e l 1 u l o s e .
L i g n i n s a r e t h e m a j o r n o n - c a r b o h y d r a t e component o f wood.
They a r e v e r y c o m p l e x , c r o s s 1i n k e d , t h r e e d imens i o n a l p o l y m e r s
formed f r o m p h e n o l i c un i t s . The number o f b u i I d i ng un i t s v a r i e s
cons i d e r a b l y f r o m a few up t o s e v e r a l h u n d r e d s . The a roma t î c na-
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t u r e o f t h e p h e n o l i c u n i t s r e n d e r s 1i g n i η h y d r o p h o b i c and t h e
Publication Date: June 1, 1977 | doi: 10.1021/bk-1977-0043.ch001
Literature Cited
Wood Anatomy
Wood Chemistry