Wood Structure and Chemical Composition

1
W o o d : Structure and Chemical Composition
R. J. THOMAS
Department of Wood and Paper Science, North Carolina State University,
Raleigh, N.C. 27607
W i t h i n living t r e e s , wood is produced to perform the r o l e s

Downloaded by 103.68.199.82 on May 4, 2018 | https://pubs.acs.org
Publication Date: June 1, 1977 | doi: 10.1021/bk-1977-0043.ch001
of support, conduction and storage. The support r o l e enables the

t r e e stem to remain e r e c t d e s p i t e the heights to which a t r e e
grows. Because of these h e i g h t s , wood a l s o must perform the r o l e
of conduct i o n , that is, transport water from the ground to the
upper parts o f the t r e e . F i n a l l y , food i s stored i n certain parts
of the wood until required by the living t r e e . The wood cells
which perform the r o l e o f conduction and/or support make up 60 to
90 percent of the wood volume. W i t h i n the living t r e e these c e l l s
are dead, that is, the cytoplasm i s absent 1eaving a hollow cell
with rigid walls. The only living cells w i t h i n the wood portion
of a t r e e are the f o o d - s t o r i n g cells. The c l o s e r e l a t i o n s h i p be
tween "form and f u n c t i o n " simplifies the study o f wood anatomy i f
the r o l e o f the cells i n the living t r e e is kept c o n s t a n t l y in
view.
Gross Anatomical Features
The e n d v i e w o f a l o g ( F i g u r e 1) e x p o s e s t h e wood and b a r k

p o r t i o n o f a t r e e t r u n k . Each y e a r a g r o w i ng c e n t e r l o c a t e d b e
tween t h e wood and b a r k i n s e r t s a new 1 a y e r o f wood a d j a c e n t t o
t h e e x i s t i ng wood. I η a d d i t i o n , new b a r k i s depos i t e d n e x t t o
t h e p r e - e x i s t i n g b a r k . Wood o c c u p i e s t h e l a r g e s t volume o f a
t r e e stem b e c a u s e more wood c e l l s a r e p r o d u c e d t h a n b a r k c e l l s
and a l s o b e c a u s e t h e wood e e l 1 s a r e r e t a i ned and t h u s a c c u m u l a t e
w h i l e t h e o u t e r m o s t b a r k e e l 1 s a r e contînually d i s c a r d e d .
The c e n t r a l wood p o r t i o n o f t h e l o g d e p i c t e d i n F i g u r e 1 i s
c o n s i d e r a b l y d a r k e r i n c o l o r than t h e p a r t a d j a c e n t t o t h e bark.
The d a r k - c o l o r e d wood i s termed heartwood and t h e 1 i g h t - c o l o r e d
wood i s termed sapwood. The d i s c o l o r a t i o n i s due t o t h e p r o d u c -
t i o n and s e c r e t i o n o f s u b s t a n c e s w h i c h a r e a b y - p r o d u c t o f t h e
death o f food-storage c e l l s . As new wood, t h a t i s sapwood, i s
formed t o t h e o u t s i d e o f t h e t r e e s t e m , add i t i o n a l i n t e r i o r s a p -
wood a d j a c e n t t o t h e h e a r t w o o d zone i s c o n v e r t e d t o h e a r t w o o d .
Some t r e e s do n o t f o r m d i s c o l o r e d h e a r t w o o d upon t h e d e a t h o f
Goldstein; Wood Technology: Chemical Aspects

ACS Symposium Series; American Chemical Society: Washington, DC, 1977.
2 WOOD TECHNOLOGY : CHEMICAL ASPECTS
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.
Figure 2. End view of a softwood showing growth rings.

Each growth ring consists of a light and dark area. The
light portion is called springwood or earlywood, and the
dark area is termed summerwood or latewood. 2 5 χ

1. THOMAS Wood: Structure and Chemical Composition 3
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 -
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 -
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

4 WOOD TECHNOLOGY: CHEMICAL ASPECTS
Figure 3. Softwood block showing

three complete and part of two other
growth rings in the cross-sectional plane
(X). Individual springwood cells can be
detected, whereas the smaller summer-
wood cells cannot be seen as individual
cells. Also note the absence of vessels

and the uniformity of the wood. Two

longitudinal surfaces (R—radial; Τ—
tangential) are illustrated. Food-storing
cells can be easily detected on the radial
surface (arrow). 47 X (Courtesy ofN.C.
Brown Center for Ultrastructural Stud
ies, S.U.N.Y. College of Environmental
Science and Forestry)
Figure 4. Cross-sectional view of springwood and sum

merwood cells. The springwood cells are larger and char
acterized by thin cell walls and a large central opening.
The smaller cells with thick cell walls and a small central
opening are summerwood cells. The narrow ribbon-like
rows of cells traversing both the springwood and sum
merwood zones (arrow) are called rays. They consist of
transversely oriented food-storage cells. 135χ

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
d i f f e r e n c e between hardwood and s o f t w o o d anatomy. Note t h e l a c k

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

Figure 5. Cross-sectional and

longitudinal surfaces of a ring-
porous hardwood. In the cross-
sectional view (X) the largest
diameter cells are springwood
vessels whereas the smaller cells

with obvious openings are sum
merwood vessels. Smaller di
ameter, thick-walled fibers con
stitute most of the remaining
volume. Transversely oriented
food-storing cells can be seen on
the radial surface (arrow). 40χ
(Courtesy of N. C. Brown Cen
ter for Ultrastructural Studies,
S.U.N J. College of Environ
mental Science and Forestry)
Figure 6. Cross-sectional and

longitudinal views of a diffuse-
porous wood. Note in cross sec
tion that the vessels, the large
diameter cells, are essentially
the same size throughout the
growth ring. Both longitudinal
views reveal the vessels (V) are
formed as the result of indi
vidual vessel cells stacked one
on top of the other in the longi
tudinal direction. The majority
of the remaining cells are small
diameter fibers. 70 χ (Courtesy
of N. C. Brown Center for Ultra-
structural Studies, S.U.N.Y. Col
lege of Environmental Science
and Forestry)

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
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
Figure 7. Isolated springwood (early-

wood) and summerwood (latewood) longi
tudinal tracheids. Note resemblance of

tracheids to long cylindrical tubes. Tracheid
lengths in this figure are considerably re
duced as tracheids are normally about 100
times longer than wide. Note the rounded
end of the springwood tracheid in the radial
view and the pointed end in the tangential
view. Summerwood tracheid ends tend to
be pointed in both views, a. Bordered pits
to adjacent longitudinal tracheids; b. and c.
pits to adjacent ray cells. (Drawing from:
Howard, E. T., Manwiller, E. G., Wood
Science (1969) 2,77-86)
Figure 8. View of internal cell walls of

springwood longitudinal tracheids. The
circular dome-like structures are bor
dered pits which permit liquid flow
between contiguous longitudinal tra
cheids. The smaller egg-shaped pits in
clusters lead to adjacent transversely
oriented ray cells. 400χ (Courtesy of
N. C. Brown Center for Ultrastructural
Studies, S.U.N.Y. College of Environ

Figure 9. Innermost wall layer as seen

from the inside of a springwood longi
tudinal tracheid showing two bordered
pits. Note the circular pit apertures and
the stringlike microfibrils which are ori
ented at approximately 90° to the long
axis of the cell. Arrow indicates longi
tudinal axis of the cell. 3,000χ
Figure 10. View of a bordered pit membrane with the

dome-shaped pit border removed. The dark central portion
is the torus. The stringlike microfibrils radiating from the
torus constitute the margo portion of the pit membrane.
Water flows freely from cell to cell through the openings
between the margo microfibrils. 3,000X

Figure 11. Springwood longitudinal tra

cheids showing bordered pits in cross-
sectional views. Note also the longitudinal
surface views of bordered pits inside of the

tracheids. In addition, narrow, elongated,
transversely oriented food-storage cells
which constitute rays are visible. 400X
(Courtesy of N. C. Brown Center for Ultra-
structural Studies, S.U.N.Y. College of
Environmental Science and Forestry)
Figure 12. Cross-sectional view of a bordered pit-pair.

A: pit aperture; B: pit border; T: torus. Note the thick,
nonperforated torus. Most of the thin margo was destroyed
during specimen preparation, and only portions of it re
main (arrow). Liquid flow occurs through the pit aper
ture, around the torus through the margo, and out the
other pit aperture into the adjacent cell. 5,000χ

a p e r t u r e o f one c e l l , t h r o u g h t h e margo p o r t i o n o f t h e membrane

and o u t t h e p i t a p e r t u r e i n t o t h e lumen o f t h e a d j a c e n t c e l l .
C r o s s - s e c t i o n a l views o f bordered p i t - p a i r s i n t e r c o n n e c t i n g t r a
c h e i d s a r e i l l u s t r a t e d i n F i g u r e 11. A v i e w o f a s i n g l e b o r d e r e d
pi t - p a i r i n F i g u r e 12 r e v e a l s an e x t r e m e l y t h i η and d i s r u p t e d
margo and a r e l a t i v e l y t h i c k t o r u s . B e c a u s e o f t h e t h i n and p o r
ous n a t u r e o f t h e margo r e g i o n , i t i s o f t e n d i s r u p t e d when p r e
p a r i n g specimens f o r m i c r o s c o p i c e x a m i n a t i o n .
Not i c e i n F i g u r e 12 t h a t t h e t o r u s i s i n t h e c e n t r a l pos i -
t i o n whîle i n F i g u r e 13 i t has moved t o one s i d e and e f f e e t i v e l y
s e a l e d one o f t h e a p e r t u r e s . I η t h i s cond î t i on t h e p i t i s i n t h e
aspirated state. C o m p a r i s o n o f an a s p i r a t e d p i t - p a i r ( F i g u r e s 13
and 1k) w i t h a n o n - a s p i r a t e d p i t - p a i r ( F i g u r e s 11 and 12) c l e a r l y
i 1 1 u s t r a t e s t h e d i s p l a c e m e n t o f t h e p i t membrane a g a i n s t t h e p i t
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
O b v i o u s l y , s o f t w o o d anatomy i s relatî v e l y s i mp1e as o n l y two

t y p e s o f e e l 1 s, l o n g i t u d i n a l t r a c h e i d s and r a y parenchyma, c o n -
s t i t u t e t h e b u l k o f t h e wood. Hardwoods have a more c o m p l e x a n a -
tomy a s more k i n d s o f e e l 1 s a r e p r e s e n t . The r o l e s o f c o n d u c t i o n
and s u p p o r t a r e c a r r i e d o u t by d i f f e r e n t c e l l s and i n add i t i o n t o
t h e t r a n s v e r s e r a y parenchyma, f o o d - s t o r a g e e e l Is o r i e n t e d i n t h e
l o n g i tud i nal d i r e c t i o n a r e p r e s e n t . Parenchyma o r i e n t e d l o n g i -
t u d i n a l l y a r e c a l l e d l o n g i t u d i n a l o r a x i a l parenchyma. Vessel
segments p e r f o r m t h e c o n d u c t i o n r o l e , and f i b e r s t h e s u p p o r t r o l e .

Figure 13. Cross-sectional view of an aspirated bordered pit-

pair. The pit membrane has moved to the border and sealed a
pit aperture with the torus. In this condition, liquid flow no
longer occurs between contiguous cells. 5,000χ
Figure 14. Surface view of an aspirated bordered pit mem

brane. The imprint of the pit aperture through the torus is the
result of an extremely tight seal. 6,200X

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 -
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

Figure 15. View from the inside of a lon

gitudinal tracheid showing pits connecting
a longitudinal tracheid to a ray cell. Note
the lack of openings within the pit mem
brane. 2,500χ
Figure 16. Types of vessel segments

found in hardwoods. A and B: springwood
vessels from a ring-porous wood. Note the
short length compared with the diameter.
C and D: typical vessel elements from dif
fuse-porous woods with simple perforation
plates at each end. E: typical diffuse-por
ous vessel element with scalariform per Ε
foration plates at each end. 1 4 0 χ

26% w h e r e a s în h i c k o r y up t o 67% o f t h e volume i s made up o f

fibers. In most c a s e s t h e h i g h e r t h e p e r c e n t a g e volume o f f i b e r s
present the higher the s p e c i f i c g r a v i t y . E x c e p t i o n s , when n o t e d ,
a r e u s u a l l y due t o t h e f i b e r s b e i n g r e l a t i v e l y t h i n - w a l l e d .
F i g u r e s 6 and 18 d e p i c t a wood i n w h i c h f i b e r s o c c u p y more t h a n
50% o f t h e vo1ume. A l s o i 1 1 u s t r a t e d a r e t h e o b v i o u s d î f f e r e n c e s
i n t h e relatî ve s i z e , shape and w a l l t h i c k n e s s between fî b e r s and
v e s s e l s.
Cons i d e r a b l e v a r i a t i o n i n t h e amount o f t r a n s v e r s e and l o n -
g i tudî n a l parenchyma e x i s t s among hardwood s p e c i e s . For example,
basswood has a p p r o x i m a t e l y t h e same as s o f t w o o d s , t h a t i s a b o u t
10%, w h i l e some oak s p e c i e s a p p r o a c h k0% parenchyma. As i n s o f t -
wood s, t h e parenchyma a r e u s u a l l y b r i c k - s h a p e d e e l 1 s a l though
some v a r i a t i o n s o f t h i s shape o c c u r s . The r a y s , composed o f

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

Figure 17. Fit membrane from

an intervessel bordered pit. Note
the absence of a torus and de-
tectable openings in the mem
brane. 2,400χ
Figure 18. Hardwood specimen

showing a vessel cell, fibers, and
ray cells. Note the retotive dif
ferences in size and shape of the
cells. The tangential view shows
a ray which is up to seven cells
wide. (Courtesy of N. C. Brown
Center for Ultrastructural Stud
ies, S.U.N.Y. College of Environ

from a s u b s t a n t i a l i n c r e a s e î n the t h i c k n e s s o f the l a y e r and

1 i t t l e o r no i n c r e a s e i n t h e S. and S. 1 a y e r s . F o r e x a m p l e , n o t e
i n F i g u r e 2 0 , w h i c h d e p i c t s a d j a c e n t v e s s e l and f i b e r c e l l w a l l s ,
t h e t h i n S 1 a y e r î η t h e t h i η v e s s e l w a l 1 and t h e t h î c k S£ i n t h e
th î c k f i b e r w a l 1 . Not i ce a l s o t h a t t h e S^ a n d S- o f b o t h e e l 1 s
a r e e s s e n t i a l l y t h e same s i z e . The a v e r a g e r e l a t i v e s i z e 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 i n d î c a t e d i n T a b l e I . The t h i n n e s t
i s o f c o u r s e t h e p r i m a r y w a l 1 w i t h t h e S~, S^ and in order of
i n c r e a s i ng t h î c k n e s s .
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
Relative Average
Thickness Angle o f
Wal1 Layer (%) MicrofibriIs
P.W. >1 Random

S 10-22 50-70°
S* 70-90 10-20°
2-8 60-90°
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 .
Chemical Compos i t i o n o f C e l l Wal1
Chemically the eel 1 wal1 i s rather heterogeneous, c o n s i s t i n g

p r i m a r i l y o f t h r e e po1yme r î c materîals: e e l 1 u l o s e , h e m i c e l l u l o s e
and 1 i g n i n. T h e s e m a t e r îals a r e composed o f l a r g e m o l e c u l e s and
c o n s t i t u t e f r o m 9 5 t o 98% o f t h e c e l l w a l l . The r e m a i n i n g 2 - 5 %
a r e l o w e r m o l e c u l a r w e i g h t compounds c a l l e d e x t r a c t i v e s . The
amount o f e a c h component, e s p e c i a l l y t h e h e m i c e l l u l o s e s , 1 i g n i n
and e x t r a c t i v e s , v a r i es c o n s î d e r a b l y between hardwoods and s o f t -
woods ( T a b l e I I ) . O t h e r f a c t o r s s u c h as s p e c i e s , l o c a t i o n o f
e e l 1 s wi t h i n t h e t r e e and g r o w t h e n v i ronment a l s o î n f 1 u e n c e t h e
f i n a l chemi c a l c o m p o s i t i o n .

Figure 19. Cell walls in cross-sectional view from contiguous

springwood longitudinal tracheids depicting wall layering. C:
compound middle lamella. 1: Sj layer; 2: S layer; and 3: S layer.
2 s
Note the S layers are the largest. 16,000 X

2
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
walled fiber and the small S layer in the thin-walled vessel.

2
13β00χ

Figure 21. Cellulose microfibrils in the primary

wall and Sj portion of the secondary wall from a
longitudinal tracheid. Note the loosely packed
and randomly arranged microfibrils in the pri-
mary wall (P). The S layer (S) consists of tightly
packed, parallel microfibrils. 12,000X

Table ! I
Average % Chemical C o m p o s i t i o n o f Softwoods and Hardwoods
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
res i dues v a r i es f r o m 7,000 t o 10,000. The c e l l u i o s e m o l e c u l e s

are 1i nked l a t e r a l l y by h y d r o g e n bonds i n t o 1 i n e a r b u n d l e s . The

e x t r e m e l y l a r g e number o f h y d r o g e n bonds r e s u l t s i n a s t r o n g l a t -
e r a l a s s o c i a t i o n o f t h e 1i n e a r c e l l u i o s e m o l e c u l e s . T h i s s t r o n g
a s s o c i a t i o n and a l m o s t p e r f e c t alîgnment o f t h e c e l 1 u l o s e mole-
c u l e s g i v e s r i s e t o c r y s t a l 1i ni t y . X - r a y measurements show t h a t
the c r y s t a l 1i ne r e g i o n s a r e i n t e r r u p t e d e v e r y 600 a n g s t r o m s wi t h
n o n - c r y s t a l 1i ne (amorphous) r e g i o n s . Whether t h i s i s due t o mi nor
i m p e r f e c t i o n s i n t h e c r y s t a l 1i ne l a t t i c e o r a r e a l s t r u c t u r a l e n -
t i t y i s not c o m p l e t e l y c l e a r . The most w i d e l y h e l d c o n c e p t i s
the 1atter i n which the c e l 1 u l o s e molecules a r e h i g h l y o r i e n t e d
( c r y s t a 1 1 i n e ) f o r a d i s t a n c e o f about 600 A, t h e n p a s s t h r o u g h an
a r e a o f poor o r i e n t a t i o n (amorphous) and r e - e n t e r a c r y s t a l 1i ne
r e g i o n . The p a t t e r n r e p e a t s t h r o u g h o u t t h e l e n g t h o f t h e c e l 1 u -
1 ose m o l e c u l e .
E l e c t r o n microscopy studies revealed t h r e a d l i k e s t r u c t u r e s
ca1 l e d microfî b r i 1 s ( F i g u r e 2 1 ) . Mi c r o f i b r i1 w i d t h s , depend î ng
upon m a t e r i a l s o u r c e and methods u s e d , v a r y f r o m 100 t o 300 ang-
s t r o m s and a r e a b o u t h a l f as t h i c k as t h e y a r e w i d e . Microfibri1
l e n g t h has n o t been d e t e r m i n e d n o r has t h e înternal s t r u c t u r e o f
the microfibrî1 been c l e a r l y e s t a b l i s h e d . Some învestigators be-
l i e v e t h a t t h e m i c r o f i b r i 1 cons i s t s o f a c r y s t a l 1i ne c o r e o f c e l -
1 u l o s e s u r r o u n d e d by an amorphous r e g i o n c o n t a i ni ng hemi c e l 1 u -
loses. Depend i ng upon t h e î n v e s t i g a t o r ^ e s t i m a t e s o f t h e c r y s -
t a l 1 i ne c o r e s i ze r a n g e s f r o m 30 by 50 A t o 40 by 100 Â. F u r t h e r
v a r i a t i o n i n t h e s i z e o f t h e microfî b r i1 i s due t o t h e amount o f
hemi c e l 1 u l o s e s s u r r o u n d i ng t h e c r y s t a l 1î ne c o r e as w e l l as a g g r e -
g a t i o n s o f s îngle m i c r o f i b r i 1 s t o f o r m m i c r o f i b r i 1 s wi t h 1 a r g e r
d iameters. S t u d i e s wi t h v e r y h i g h r e s o l u t i o n e l e c t r o n m i c r o s c o p e s
c o u p l e d wi t h n e g a t i ve s t a î ni ng t e c h n i q u e s has l e d t o t h e v i e w
t h a t a 35 Â w i d e c e l 1 u l o s e f i b r i l c a l l e d t h e e l e m e n t a r y f i b r i l i s
the bas î c s t r u c t u r e . One p r o p o s a l f r o m t h i s work v i e w s t h e m i c r o -
f i b r i l as cons i s t i ng o f f o u r e l e m e n t a r y f i b r i l s wi t h h e m i c e l 1 u -
l o s e s , l i g n i n and w a t e r a r o u n d t h e o u t s i d e as w e l l as between t h e
elementary f i b r i I s . More r e c e n t w o r k has i n d i c a t e d s o - c a l l e d
s u b - e l e m e n t a r y f i b r i I s i n t h e 10 A r a n g e . O b v i o u s l y t h e phys i c a l
s t r u c t u r e o f c e l 1 u l o s e i n t h e woody pi a n t c e l 1 w a l 1 i s f a r f r o m

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-
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
t h r e e d imens i o n a l n e t w o r k p r o v i d e s r i g i d î t y and o p t îmum r e s i s t a n c e

to compressive f o r c e s . The s t r u c t u r a l make-up o f 1 i g n i n f r o m
hardwoods d i f f e r s f r o m s o f t w o o d 1 i g n i n î η t h a t t h e bas i c bu i I d i ng
u n i t i s s y r i n g y l w h e r e a s gua i a c y 1 i s t h e p r imary bu îId i ng un i t
f o r softwoods. B e c a u s e t h e gua i a c y 1 u n i t has a g r e a t e r number o f
p o t e n t i a l r e a c t i ve s i t e s , a h i gher d e g r e e o f c r o s s l i n k i n g e x i s t s .
F u r t h e r m o r e 1 i g n i η formed pr i marîly f r o m gua i a c y 1 h a s , on t h e
a v e r a g e , a h i g h e r m o l e c u l a r we î g h t . T y p i c a l l y , s o f t w o o d 1i g n i η
has a b o u t t e n t i m e s more g u a i a c y l t h a n s y r i n g y l un i t s w h e r e a s i n
hardwoods t h e r a t i o i s u s u a l l y one t o one. Thus hardwood 1 i g n i n s
a r e more eas i l y d e g r a d e d t h a n s o f t w o o d 1i gn i n s .
The d i s t r i but i o n o f t h e c h e m i c a l c o n s t i t u e n t s a c r o s s t h e
c e l 1 wal1 i s not un i f o r m . Wi t h r e g a r d t o c e l 1 u l o s e , t h e p r i m a r y
wal1 c o n t a î ns a b o u t 10% i n c r e a s i ng t o more t h a n 50% i n t h e lay-
e r and d e c r e a s i ng s i i g h t l y i n t h e S 1 a y e r . The 1 i g n i n c o n t e n t
o f t h e m i d d l e 1ame11 a and p r i m a r y wal1 i s on t h e o r d e r o f 70%.
The S c o n t a i n s a b o u t 22% and t h e S a p p r o x i m a t e l y 15%. I t should
be n o t e d t h a t due t o t h e v e r y 1arge volume o f t h e S l a y e r , over
h a l f o f t h e t o t a l 1 i g n i n i s found i n t h î s l a y e r a l t h o u g h t h e c o n -
centrât i o n i s l o w e r t h a n i n t h e compound m i d d l e 1 a m e l i a . A l s o ,
b e c a u s e t h e compound m i d d l e l a m e l l a i s so t h î η o n l y 10% o f t h e
t o t a l 1i g n i n i s p r e s e n t d e s p i t e t h e v e r y h i g h c o n c e n t r a t i o n . The
h e m i c e l 1 u l o s e f r a c t i o n t e n d s t o v a r y a b o u t t h e same as t h e c e l 1 u -
l o s e a c r o s s the c e l 1 wal1.
I η add i t i o n t o t h e m a j o r c e l 1 wal1 components o f c e l 1 u l o s e ,
h e m i c e l 1 u l o s e and 1 i g n i n, wood c o n t a i n s v a r y i ng amounts o f sub
s t a n c e s termed extractî v e s . The t e r m e x t r a c t i ves i n c l u d e s a w i d e
range o f chemi c a l t y p e s and a v e r y 1arge number o f i nd i v i d u a l
compounds. Some o f t h e m a j o r chemi c a l t y p e s a r e 1) T e r p e n e s and
r e l a t e d compounds, 2) F a t t y a c i d s , 3) A r o m a t i c compounds and 4)
Volâtile o i l s . S p e c i e s d i f f e r w i d e l y i n t h e t y p e and amount o f
extractî ves p r e s e n t . A l s o t h e r e i s cons i d e r a b l e v a r i a t i o n i n t h e
d i s t r i b u t i o n o f e x t r a c t îves t h r o u g h o u t t h e wood o f i n d i v i d u a l
trees. A l t h o u g h some e x t r a c t i v e s a r e f o u n d i n sapwood, t h e h e a r t -
wood u s u a l l y c o n t a i ns t h e l a r g e s t amount.
A l t h o u g h t h e e x a c t c a u s e o f h e a r t w o o d f o r m a t i o n i s not known,

the r e s u l t i ng changes have been w e l l d o c u m e n t e d .The parenchyma

eel 1 s change thei r metabolic a c t i v i ty and tend to produce e x t r a c -
t i v e s from stored c a r b o h y d r a t e s . The parenchyma cel1 s d i e and
the e x t r a c t ives d i ffuse i n t o adjacent cel1 s . Thus, the heartwood
conta i ns no 1i v i ng cel1 s and in many species is d î s c o l o r e d as a
r e s u l t o f the e x t r a c t i ve c o n t e n t . I η c e r t a i η softwoods res in
canal s are another source o f e x t r a c t i v e s . These s t r u c t u r e s con
ta i η cons i d e r a b l e quanti t i e s o f res in in the 1 î v i ng t r e e .
Despi te the r e l a t i v e l y low percentage content o f e x t r a c t î ves
(Table I I ) , they very often i nf1uence wood propert ies and thus
play a r o l e i n u t i 1 i z a t i o n . Advantages accrue from the presence
of c o l o r e d and vol at i l e e x t r a c t ives which provide e s t h e t i c values.
Some of the phenolic compounds provide r e s i s t a n c e to i n s e c t and
fungal a t t a c k . Other e x t r a c t i ves provî de useful products. From
ta 11 o i 1 , products such as turpent i ne, ros in and f a t t y a c i d s are

p r o d u c e d .I η add i t i o n , tanni ns, camphor, gum a r a b i c , n a t u r a l
rubber and f1avonoids are some of the many products from e x t r a c -
t i ves.
E x t r a c t i ves somet imes prevent or i nhi bi t the ut î1î zat ion o f
w o o d . For example, woods contai ni ng phenolic type e x t r a c t i ves
cannot be pulped v i a the s u l f î te p r o c e s s . The s o - c a l l e d " p i t c h
t r o u b l e s " in the p u l ρ and paper î n d u s t r y r e s u l t from the tendency
of the res i η type e x t r a c t i ves to coagulate and adhere to metal
and f i b r o u s s u r f a c e s . A l s o the presence of e x t r a c t i v e s r e s u l t
in a h i gher consumpt ion of pulpi ng chemical s and i n lower p u l ρ
yields.
Despi te the numerous products d e r i ved from e x t r a c t i v e s , much
of the bas i c chemi s t r y o f the numerous compounds found in many
species i s s t i l l v i r t u a l l y u n k n o w n .Future work in th i s area
should provî de many new products.
Literature Cited
Wood Anatomy
1. I senberg, I. H. "The S t r u c t u r e of Wood i n The Chemistry o f

Wood" e d i t e d by B. L. Browning. I n t e r s c i e n c e P u b l i s h e r s ,
John Wiley and Sons. New York, New York. 1963.
2. Jane, F. W. "The S t r u c t u r e o f Wood" 2nd Edition. Adam and
Charles Black. London, England. 1970.
3. Panshin, A . G. and Carl d e Z e e u w ."Textbook o f Wood Technol-
ogy" 3rd Edition. McGraw-Hill. New York, New York. 1970.
Wood Chemistry
1. Browning, B. L. Editor. "The Chemistry o f Wood" I n t e r -

science P u b l i s h e r s , John Wiley and Sons. New York, New York.
1963.
2. Hanna, R. B. and W. A . Côté, Jr. The Sub-elementary Fibril
of P l a n t Cell Wall Cellulose. C y t o b i o l o g i e (1974) 10(1):102.

3. Hillis, Ε. W. "Wood Extractives and Their Significance to

t h e P u l p and P a p e r I n d u s t r y " A c a d e m i c Press. New Y o r k , New
York. 1962.
4. P r e s t o n , R. D. "The Physical B i o l o g y o f Plant Cell W a l l s "
Chapman and Hall. London. 1974.
5. W i s e , L. E. and E. C. Jahn. "Wood C h e m i s t r y Volume I " ACS
Monogram Series. Reinhold Publishing Corporation. New Y o r k ,
New Y o r k . 1 9 5 2 .


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1

W o o d : Structure and Chemical Composition

W i t h i n living t r e e s , wood is produced to perform the r o l e s

of support, conduction and storage. The support r o l e enables the

Gross Anatomical Features

The e n d v i e w o f a l o g ( F i g u r e 1) e x p o s e s t h e wood and b a r k

Goldstein; Wood Technology: Chemical Aspects

Figure 2. End view of a softwood showing growth rings.

Goldstein; Wood Technology: Chemical Aspects

Goldstein; Wood Technology: Chemical Aspects

Figure 3. Softwood block showing

cells. Also note the absence of vessels

and the uniformity of the wood. Two

Figure 4. Cross-sectional view of springwood and sum­

Goldstein; Wood Technology: Chemical Aspects

d i f f e r e n c e between hardwood and s o f t w o o d anatomy. Note t h e l a c k

Goldstein; Wood Technology: Chemical Aspects

Figure 5. Cross-sectional and

vessels whereas the smaller cells

Figure 6. Cross-sectional and

Goldstein; Wood Technology: Chemical Aspects

Goldstein; Wood Technology: Chemical Aspects

Figure 7. Isolated springwood (early-

tudinal tracheids. Note resemblance of

Figure 8. View of internal cell walls of

Goldstein; Wood Technology: Chemical Aspects

Figure 9. Innermost wall layer as seen

Figure 10. View of a bordered pit membrane with the

Goldstein; Wood Technology: Chemical Aspects

Figure 11. Springwood longitudinal tra­

surface views of bordered pits inside of the

Figure 12. Cross-sectional view of a bordered pit-pair.

Goldstein; Wood Technology: Chemical Aspects

a p e r t u r e o f one c e l l , t h r o u g h t h e margo p o r t i o n o f t h e membrane

O b v i o u s l y , s o f t w o o d anatomy i s relatî v e l y s i mp1e as o n l y two

Goldstein; Wood Technology: Chemical Aspects

Figure 13. Cross-sectional view of an aspirated bordered pit-

Figure 14. Surface view of an aspirated bordered pit mem­

Goldstein; Wood Technology: Chemical Aspects

Goldstein; Wood Technology: Chemical Aspects

Figure 15. View from the inside of a lon­

Figure 16. Types of vessel segments

Goldstein; Wood Technology: Chemical Aspects

26% w h e r e a s în h i c k o r y up t o 67% o f t h e volume i s made up o f

some v a r i a t i o n s o f t h i s shape o c c u r s . The r a y s , composed o f

Goldstein; Wood Technology: Chemical Aspects

Figure 17. Fit membrane from

Figure 18. Hardwood specimen

Goldstein; Wood Technology: Chemical Aspects

from a s u b s t a n t i a l i n c r e a s e î n the t h i c k n e s s o f the l a y e r and

P.W. >1 Random

Chemical Compos i t i o n o f C e l l Wal1

Chemically the eel 1 wal1 i s rather heterogeneous, c o n s i s t i n g

Goldstein; Wood Technology: Chemical Aspects

Figure 19. Cell walls in cross-sectional view from contiguous

Note the S layers are the largest. 16,000 X

walled fiber and the small S layer in the thin-walled vessel.

Goldstein; Wood Technology: Chemical Aspects

Figure 21. Cellulose microfibrils in the primary

Goldstein; Wood Technology: Chemical Aspects

Average % Chemical C o m p o s i t i o n o f Softwoods and Hardwoods

res i dues v a r i es f r o m 7,000 t o 10,000. The c e l l u i o s e m o l e c u l e s

are 1i nked l a t e r a l l y by h y d r o g e n bonds i n t o 1 i n e a r b u n d l e s . The

Goldstein; Wood Technology: Chemical Aspects

Figure 4. Cross-sectional view of springwood and sum

Figure 11. Springwood longitudinal tra

Figure 14. Surface view of an aspirated bordered pit mem

Figure 15. View from the inside of a lon