4 P a r t O n e - S e c t i o n 1 B a m b o o M o r p h o l o g y a n d P h y s i o l o g y

B A M B O O M O R P H O L O G Y

A N D P H Y S I O L O G Y

P A R T S O F T H E P L A N T

1 . - T H E R H I Z O M E

B a m b o o s a r e p e r e n n i a l p l a n t s c o n s i s t i n g o f a r a m i f y i n g

s y s t e m o f s e g m e n t e d v e g e t a t i v e a x e s w h i c h f o r m a r e g u l a r

a l t e r n a t i o n o f n o d e s a n d i n t e r n o d e s . T h e s e s e g m e n t e d a x e s ,

a c c o r d i n g t o t h e i r s h a p e a n d p o s i t i o n o n t h e p l a n t , h a v e t h e

f o l l o w i n g n a m e s z t h e c u l m o r s t e m w i t h t h e b r a n c h e s , w h i c h

f o r m s t h e a e r i a l p a r t o f t h e p l a n t a n d t h e r i z o m e s y s t e m

w i t h i t s r o o t s w h i c h f o r m s t h e s u b t e r r a n e a n p a r t o f t h e

p l a n t ; a n d i t s s t r u c t u r a l f o u n d a t i o n . T h e r h i z o m e p e r f o r m s

i m p o r t a n t f u n c t i o n s i n t h e l i f e o f t h e p l a n t ; i t i s t h e v i t a l

o r g a n w i t h w h i c h b a m b o o p l a n t s r e p r o d u c e v e g e t a t i v e l y o r

a s e x u a l l y t h r o u g h t h e b r a n c h i n g o f r h i z o m e s . A s a n o r g a n ,

i t h a s t h e f u n c t i o n o f s t o r a g e a n d t r a n s p o r t a t i o n o f n u t r i -

e n t s . T h e c u l m s d e p e n d o n r h i z o m e s f o r t h e i r g r o w t h , v i g o r

a n d s p a c i n g o n t h e g r o u n d .

T y p e s o f b a m b o o s

A s m e n t i o n e d a b o v e , t h e t r e e s p e c i e s a r e c o m m o n l y

d i v i d e d i n t o t w o l a r g e g r o u p s a c c o r d i n g t o t h e i r g e o g r a p h i -

c a l p o s i t i o n . T h e y a r e : " s o f t w o o d s " w h i c h n a t u r a l l y g r o w i n

t h e t e m p e r a t e z o n e s , a n d h a r d w o o d s w h i c h g r o w i n t h e

t r o p i c a l z o n e s . T h e s e t w o g r o u p s h a v e d i f f e r e n c e s p a r t i c u -

l a r l y i n t h e a n a t o m y o f t h e i r t r u n k s a n d t h e i r l e a v e s .

L i k e t r e e s , b a m b o o s a r e a l s o c l a s s i f i e d i n t w o m a i n

t y p e s o n t h e f o r m a t i o n p a t t e r n o f t h e s u b t e r r a n e a n p a r t o f

t h e c u l m w h i c h a l s o d e p e n d s o n t h e g e o g r a p h i c a l p o s i t i o n

w h e r e t h e y g r o w s n a t u r a l l y . T h e s e m a i n g r o u p s o r t y p e s

a r e : t h e l e p t o m o r p h o r m o n o p o d i a l t y p e w h i c h g r o w s i n

t e m p e r a t e z o n e s ; a n d t h e p a c h y m o r p h o r s y m p o d i a l t y p e

w h i c h g r o w s i n t h e t r o p i c a l z o n e s . T h e r e i s a s u b g r o u p o r

i n t e r m e d i a t e k n o w n a s m e t a m o r p h w h i c h i s a c o m b i n a t i o n

o f t h e t w o m a i n g r o u p s , b u t h a s n o r e l a t i o n w i t h t h e g e o -

g r a p h i c a l p o s i t i o n .

T h e o n l y t h i n g t h a t t h e s e t w o g r o u p s h a v e i n c o m m o n

i s t h e m o r p h o l o g y a n d a n a t o m y o f t h e i r c u l m s a n d t h e i r

g r o w i n g p r o c e s s . T h e a e r i a l p a r t o f t h e s e t w o g r o u p s i s s o

s i m i l a r t h a t i f t h e s p e c i e s a r e n o t k n o w n , i t i s d i f f i c u l t t o

r e c o g n i z e v i s u a l l y t o w h i c h g r o u p t h e y b e l o n g , u n l e s s t h e

r h y z o m e s c a n b e s e e n , b e c a u s e t h e r e i s a g r e a t d i f f e r e n c e

i n t h e m o r p h o l o g y a n d i n t h e f o r m o f b r a n c h i n g o f t h e r h i -

z o m e s b e t w e e n t h e t w o m a i n g r o u p s a n d i n t h e f o r m a t i o n

o f t h e c u l m s h o o t , a s c a n b e s e e n i n t h e F i g 1 . 1 .

O n t h e o t h e r h a n d , t h e t w o m a i n g r o u p s o f b a m b o o s

h a v e d i f f e r e n c e s i n t h e n u m b e r o f c h r o m o s o m e s , w h i c h i n

t h e l e p t o m o r p h t y p e i s 2 n = 4 8 , a n d i n t h e p a c h y m o r p h t y p e

i s 2 n = 7 2 . T h e b a s i c n u m b e r o f c h r o m o s o m e s i s a s s u m e d t o

b e 2 n = 1 8 . B a m b o o s w i t h l o w m u l t i p l e c h r o m o s o m e s a r e

c o n s i d e r e d t o b e o f t h e a d v a n c e d t y p e a n d t h o s e w i t h h i g h

o n e s a r e o f p r i m i t i v e t y p e . T h e b a s i c n u m b e r f o r m o s t

b a m b o o s p e c i e s i s X = 1 2 , e x c e p t f o r c e r t a i n b a m b o o s p e c i e s

( e s p e c i a l l y p a q u i m o r p h b a m b o o s ) . T h e d i f f e r e n c e i n

c h r o m o s o m e n u m b e r a n d n u c l e o u s t y p e c a n h e l p t o

e x p l a i n t h e s y s t e m a t i c d e v e l o p m e n t , a n d c l a s i f i c a t i o n o f

b a m b o o s .

I n 1 8 7 9 , t h e R i v i e r e s w e r e t h e r s t t o p u b l i s h a c l e a r

d i s t i n c t i o n b e t w e e n t h e t w o b a s i c f o r m s o f t h e b a m b o o r h i -

z o m e s . T h e y u s e d t h e t e r m s " c a e s p i t o s e " , o r c l u m p i n g f o r

p a c h y m o r p h , a n d ' t r a c a n t " o r r u n n i n g f o r l e p t o m o r p h . I n

1 9 2 5 M c C l u r e i n t r o d u c e d t h e t e r m s m o n o p o d i a l a n d

s y m p o d i a l w h e n h e w a s l i v i n g i n C h i n a , a n d l a t e r , a t t h e

S m i t h s o n i a n I n s t i t u t i o n , i n W a s h i n g t o n , h e d e v e l o p e d t h e

c o n c e p t o f l e p t o m o r p h a n d p a c h y m o r p h r e s p e c t i v e l y .

T o d a y t h e t e r m s l e p t o m o r p h a n d p a q u i m o r p h a r e u s e d

i n t h e A m e r i c a s b y t h e b o t a n i s t s a n d t a x o n o m i s t s , a n d t h e

t e r m s m o n o p o d i a l a n d s y m p o d i a l i n A s i a .

a ) . - L E P T O M O R P H R H I Z O M E

T h e l e p t o m o r p h r h y z o m e i s a l s o k n o w n b y t h e n a m e s

o f m o n o p o d i a l , t r a c a n t , i n d e t e r m i n a t e , a n d r u n n i g b a m -

b o o s . B a m b o o s w i t h l e p t o m o r p h r h i z o m e s a r e u s u a l l y d i s -

t r i b u t e d i n t e m p e r a t e r e g i o n s , s u c h a s J a p a n , K o r e a a n d

C h i n a , w h e r e w i n t e r s a r e s e v e r e . T h e y a r e c h a r a c t e r i z e d b y

r e l a t i v e l y s t r o n g f r o s t - r e s i s t a n c e , a n d c o n s e q u e n t l y t h e y

c a n b e c u l t i v a t e d a t h i g h e l e v a t i o n s i n t h e t r o p i c s .

B a m b o o s w i t h t h i s t y p e o f r h i z o m e a r e r e p r e s e n t e d i n

A s i a b y t h e g e n e r a A r u n d i n a r i a , a n d P h y l l o s t a c h y s . T h e

m o s t c o l d - r e s i s t a n t a r e : P h y l l o s t a c h y s p r a e c o x , P h . p r o p i t i -

q u a , P h . d u l c i s , P h . i r i d e n s c e n s , P h . n u d a , P h . a n g u s t a , P h .

a u r e o s u l c a t a , a n d I n d o c a l a m u s . I n t h e A m e r i c a s t h e r e a r e

o n l y t h r e e n a t i v e s p e c i e s o f l e p t o m o r p h t y p e w h i c h b e l o n g

t o t h e a s i a t i c g e n u s A r u n d i n a r i a , a n d g r o w i n t h e t e m p e r a t e

z o n e s o f t h e s o u t h e a s t e r n o f t h e U n i t e d S t a t e s u p t o 4 6

n o r t h l a t i t u d .

T h e s u b t e r r a n e a n p a r t o f t h e p l a n t c o n s i s t s o f t w o

m a j o r p a r t s , t h e c u l m - b a s e w i t h i t s r o o t s y s t e m a n d t h e

r h i z o m e s y s t e m ( S e e F i g . 1 . 1 ) . T h e c u l m - b a s e c o r r e s p o n d s

t o t h e s u b t e r r a n e a n p r o l o n g a t i o n o f t h e a e r i a l c u l m a n d i s

c o n e c t e d t o t h e r h i z o m e b y t h e c u l m n e c k . T h e i n t e r n o d e s

a t t h e l o w e r p a r t o f t h e c u l m b a s e a r e t h e s h o r t e s t a n d t h e r e

a r e u s u a l l y 1 3 - 1 6 .

T h e l e p t o m o r p h r h i z o m e h a s t h e c r e e p i n g h a b i t . I t i s

l o n g a n d s l e n d e r , w i t h a c y l i n d r i c a l o r s u b c y l i n d r i c a l f o r m ,

a n d a d i a m e t e r u s u a l l y l e s s t h a n t h a t o f t h e c u l m s o r i g i n a t -

i n g f r o m i t . T h e i n t e r n o d e s a r e l o n g e r t h a n b r o a d , a n d t h e y

a r e g e n e r a l l y s h o r t e r t h a n t h o s e o f t h e c u l m . T h e y a r e t y p i -

c a l l y s o l i d w i t h n a r r o w l u m e n . E v e r y n o d e b e a r s a s o l i t a r y

l a t e r a l b u d a n d a n e n c i r c l i n g b e l t o f r o o t s a t t h e n o d e .

B e y o n d t h e b u d t h e r e i s a l o n g i t u d i n a l g r o o v e .

M o s t o f t h e l a t e r a l b u d s a r e t e m p o r a r i l y o r p e r m a n e n -

t l y d o r m a n t . T h e m a j o r i t y o f t h o s e t h a t g e r m i n a t e g e n e r a t e

s i n g l e c u l m s , d i r e c t l y , o r r h i z o m e s . B u t i t i s v e r y d i f f i c u l t t o

Part One- Section 1- Bamboo Morphology and Physiology 5

determine whether a lateral bud of the rhizome is going toform a rhizome or a culm shoot when it is still dormant. Themeristem generated in the lateral bud forms a culm neckwhich turns its apex upward forming the culm base and theculm shoot, and finally the culm proper. The buds on theculmbase can only grow into rhizomes and cannot growinto new culms.

The culms and the rhizomes grow in alternation allthrough the year. According to Ueda (1960), in temperatezones the rhizome grows during summer and autum whenthe temperature is relatively high, and the culms growthrough winter to spring when the temperature is relativelylow. The rhizome begins to develop after the new culm sho-ots have attained their full growth and the new branchesand leaves have developed. This usually starts in Marchafter the soil temperature rises up to about + 5 C. The rhy-zome grows fastest in August and September, and ceases togrow gradually from November onwards.

The rhizome shoot is slender and runs its apical meris-tem forward, parallel to the ground surface. The elongationof a leptomorph rhizome shoot depends on the activity oftwo meristems: the apical meristem at the terminal end ofthe rhizome, and the intercalary or intermediate meris-tems located between the internodes of the rhizome. Theapical meristem consists of tunicacorpus and its derivedmeristem. The cells of a derived meristem differenciate intosheaths, buds, primordial roots and vascular bundles. Theoriginal slanted bundles and enlarged parenchyma cells

form a new node at the young sheath position, and dividethe derived meristem into several intercalary meristems.

The growth of a rhizome is mainly determined by thestretching action of a terminal portion of the rhizome whichconsists of 14-16 actively elongating internodes. The intern-odes of the streching portion elongate according to therhythm of "slow-fast-slow" from back to front. The lattersections of the stretching portion mature and stop stret-ching. As the growing season proceeds, the apical meristemproduces new nodes and internodes continnuously. Thenew internodes formed by the meristem at the terminalend, act in succession replacing the former part of thestretching portion. The rhizome tip continues to grow.

The apical meristem of the rhizome shoot is tightlyembraced layer by layer by hard thick sheaths and issharply pointed like a borer which can penetrate throughthe hard-textured soils or the gravel gaps with driven forcegenerated from the internodal elongation. The rhizomeshoot does not necessarily maintain a horizontal positionnor does it fallows in a fixed direction but it may twist, bed,shrink, elongate or wind with the topogragraphical and soilconditions. It grows in all directions, and forms a complexinterwined network. It may run to considerables distancesand send out many single culms that appear on the groundscattered at certain distances every year.

Ueda (1960) made extensive excavations of the lepto-morph rhizomes of several species of Japanese bamboos,

Fig. 1.2 RHIZOME SYSTEM OF THE LEPTOMORPH TYPE

Cu|m ii 9 Rhizome roots The shoot generated

I it

-;~~4tifiiiittissssasg

Culm shootRhizome-

Culmbaseroots"W;-,,..nw

, , 1 Culm base

\ Culm neck~11 xv s __ ,1 , .1 Pg - ,, I

_.__. r

"'1' -i"'fQ'fs . 1 .

by the rhizome budcan form a new culm ora new rhizome

9 I*1? 9

I z _.1i

.3;-;_.-gzbyFioots

Apicalmeristem Growing rhizome-The growth of a rhizome is

mainly determined by the stretching action oi aterminal portion of the rhizome which consists of14-16 actively elongating internodes

~ - b . - . - ~ v - 4

- U .

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6 P a r t O n e - S e c t i o n 1 - B a m b o o M o r p h o l o g y a n d P h y s i o l o g y

F i g . 1 . 3 T h e s o i l h a s b e e n r e m o v e d b y U e d a i n o r d e r t o

s h o w t h e i n t r i n c a t e s y s t e m o f r h i z o m e s w h i c h f o r m a n e t -

w o r k j u s t b e l o w t h e s u r f a c e a n d c o n e c t m o s t o f t h e s t a n -

d i n g c u l m s . T h i s m e a n s t h a t t h e w h o l e g r o v e i s o n e p l a n t .

p l o t t i n g t h e p a t t e r n a n d r a t e o f t h e i r e x t e n s i o n . H e f o u n d

t h a t t h e t o t a l l e n g t h o f l i v i n g r h i z o m e s p e r 0 . 1 h a . i n b a m -

b o o g r o v e s o f P h y l l o s t a c h i s r e t i c u l a t a w a s f r o m 6 , 3 0 0 t o

1 8 . 7 4 0 m e t e r s ; i n g r o v e s o f P l e i o b l a s t u s p u b e s c e n s f r o m

4 7 , 0 0 0 t o 5 7 , 9 2 0 m e t e r s .

T h e r h i z o m e n e t h u n d e r t h e g r o u n d p r e v e n t s l a n d s l i d e

a n d c o l l a p s e o f r i v e r b a n k a n d h i l l s i d e s w h e r e t h e r e a r e a n

e r o s i o n p r o b l e m s . B a m b o o c u l t i v a t i o n f o r t h e p r o t e c t i o n o f

r i v e r b a n k s w a s r e c o m m e n d e d i n J a p a n i n t h e s i x t e e n c e n -

t u r y . T h i s i s a l s o t h e r e a s o n w h y a b a m b o o p l a n t a t i o n i s

c o n s i d e r e d a s a f e r e f u g e w h e n a n e a r t h q u a k e t a k e s p l a c e .

T h e d e p t h t h a t t h e r h i z o m e t r a v e l s o n t h e g r o u n d a l s o

v a r i e s w i t h t h e t y p e o f b a m b o o . T h e h o r i z o n t a l g r o w t h

m o s t l y t a k e p l a c e i n t h e u p p e r s o i l l a y e r b e t w e e n 1 0 - 3 0 c m

i n d e p t h , w h e r e w a t e r , h e a t , a n d a i r a r e e a s i l y a v a i l a b l e .

V e r y f e w r h i z o m e s g r o w d e e p e r t h a n 5 0 c m .

T h e a n n u a l e x t e n t i o n o f t h e r h i z o m e s r a n g e s f r o m 0 . 3 0

t o 0 . 7 0 m e t e r s i n s m a l l b a m b o o s a n d a b o u t 4 - 6 m e t e r s i n

l a r g e c u l m b a m b o o s s u c h a s P h y l l o s t a c h y s b a m b u s o i d e s

( M a d a k e ) . T h e i r l e n g t h v a r i e s a c c o r d i n g t o s p e c i e s a n d h a b i -

t a t . T h e g r o w t h p e r i o d i s s h o r t e r i n a r e a s o f h i g h l a t i t u d a n d

h i g h e l e v a t i o n s t h a n i n l o w l a t i t u d e s a n d l o w e l e v a t i o n s . T h e

s a m e i s t h e c a s e o n s h a d e d s l o p e s c o m p a r e d t o s u n n y

s l o p e s .

A c c o r d i n g t o T a k e n o u c h i ( 1 9 3 2 ) , i n s p e c i a l c o n d i t i o n s ,

t h e r h i z o m e t o p g o e s o u t o f t h e e a r t h o r s o i l a n d s o m e c o n -

t i n u e t o g r o w u p w a r d s t o b e c o m e b a m b o o s h o o t s . S o m e ,

h o w e v e r , e n t e r t h e e a r t h a g a i n a f t e r s t r e t c h i n g a b i t o n

e a r t h , f o r m i n g t h e s o c a l l e d j u m p i n g r h i z o m e " l i k e a b o w .

J u m p i n g r h i z o m e s t e n d t o a p p e a r w h e r e t h e u p p e r l a y e r o f

f o r e s t s o i l i s h a r d e n e d a n d i m p e r v i o u s . T h e s a m e a u t h o r

p o i n t s o u t t h a t w h e n t h e r h i z o m e i s i n j u r e d o r c u t o f f , t h e

n e a r e s t b u d b e h i n d t h e i n j u r e d p a r t d e v e l o p s i n t o a n e w

r h i z o m e , w h i c h c o n t i n u e s t o g r o w i n t h e s a m e d i r e c t i o n o f .

t h e p a r e n t r h i z o m e .

T h e l o n g e v i t y o f t h e r h i z o m e a l s o v a r i e s w i t h t h e s p e c i e

o f b a m b o o . A c c o r d i n g t o T a k e n o u c h i ( 1 9 3 2 ) , t h e r h i z o m e o f

P l e i o b l a s t u s s i m o n i ( M a d a k e ) a n d P h y l l o s t a c h y s n i g r a

h e n o n i s ( H a c h i c u ) c o n t i n u e t o f u n c t i o n a c t i v e l y t o t h e t h i r d

y e a r , a f t e r w h i c h t h e y g r a d u a l l y d e c l i n e i n v i g o r a n d f r o m

a b o u t t h e f i f t h y e a r t h e y s l o w l y g o i n t o d e c a y a n d d i e i n t h e

s e v e n t h o r e i g h t h y e a r . I n P . e d u l i s ( M o s o c h i c u ) t h e p e r i o d

o f h i g h e s t a c t i v i t y o f t h e r h i z o m e i s f r o m t h e t h i r d t o t h e

s i x t h y e a r . I n t h e e i g h t h o r n i n t h y e a r d e c a y s e t s i n , a n d

k i l l s i t i n t h e t w e l f t h o r t h i r t e e n t h y e a r ( T a k e n o u c h i 1 9 3 9 ,

W e n e t a l l . 1 9 8 1 . ; X i a o ? 1 9 9 1 ) .

D i r e c t i o n w h i c h t h e l e p t o m o r p h

r h i z o m e t a k e s w h e n i t s t a r t s i t s g r o w t h

T h e f o l l o w i n g i n f o r m a t i o n w a s f o u n d i n t h e C h i n e s e

b o o k T r e a t i s e o n H u s b a n d r y ( C h i m i n y a o s h u ) , w r i t t e n b y

C h i a S u n i u i n t h e 5 t h c e n t u r y : O w i n g t o t h e n a t u r e o f t h e

b a m b o o i t h a s a t e n d e n c y t o s p r e a d i t s r h i z o m e g r o w t h i n

a s o u t h w e s t e r l y d i r e c t i o n . T h e r e f o r e t h e y a r e u s u a l l y

p l a n t e d i n t h e n o r t h e a s t e r n p a r t o f t h e g a r d e n o r g r o v e .

A f t e r s e v e r a l y e a r s t h e y w i l l s p r e a d i n t h e i r g r o w t h u n t i l

t h e c u l m s f i l l t h e w h o l e g a r d e n . T h e r e i s a t r a d i d i t i o n t h a t

i f t h e b a m b o o s a r e p l a n t e d o n t h e w e s t e r n s i d e o f t h e h o u s e

t h e i r r o o t s w i l l s p r e a d , c o v e r i n g t h e g r o u n d u n t i l t h e y

r e a c h a n e i g h b o r s h o u s e o n t h e w e s t .

T a k i n g i n t o a c c o u n t t h a t C h i n a i s l o c a t e d i n t h e n o r -

t h e r n h e m i s p h e r e , I t h i n k t h a t i t i s v e r y i m p o r t a n t t o

e x p e r i m e n t t h i s i n t e r e s t i n g t h e o r y s o u t h o f t h e E c u a d o r , (

i n t h e s o u t h e r n h e m i s p h e r e ) , i n o r d e r t o s e e i f t h e r e i s a n y

c h a n g e i n t h e d i r e c t i o n t h e r h i z o m e t a k e s w h e n i t s t a r t s

i t s g r o w i n g p r o c e s s .

b ) . - P A C H Y M O R P H R H I Z O M E

T h i s t y p e o f r h i z o m e i s a l s o k n o w n a s c l u m p t y p e ,

s y m p o d i a l , c a e s p i t o s e a n d d e t e r m i n a t e . I t i s t y p i c a l o f

t r o p i c a l z o n e s o f t h e A m e r i c a s , A s i a , A f r i c a a n d O c e a n i a .

I t c a n n o t w i t h s t a n d f r e e z i n g t e m p e r a t u r e s . B a m b o o s

w i t h t h i s t y p e o f r h i z o m e a r e r e p r e s e n t e d i n t h e A m e r i c -

a s b y g e n u s G u a d u a , a n d i n t r o p i c a l A s i a b y t h e g e n e r a

D e n d r o c a l a m u s a n d B a m b u s a .

T h e m o r f o l o g y a n d g r o w i n g p r o c e s s o f p a c h y m o r p h r h i -

z o m e s i s d i f f e r e n t f r o m t h a t o f t h e l e p t o m o r p h r h i z o m e s

a n d h a s t h e f o l l o w i n g c h a r a c t e r i s t i c s : T h e c u l m - b a s e d o e s

n o t e x i s t i n t h e s u b t e r r a n e a n p a r t o f t h e c u l m a n d t h e a e r i a l

c u l m i s g e n e r a t e d d i r e c t l y b y t h e r h i z o m e , w h i c h , i n t h i s

c a s e c o u l d b e c o n s i d e r e d a s t h e c u l m - b a s e .

T h e r h i z o m e i s s o l i d , w i t h r o o t s o n t h e l o w e r s i d e , t h e

s h a p e i s u s u a l l y m o r e o r l e s s c u r v e d , a n d t h e i n t e r n o d e s a r e

b r o a d e r t h a n l o n g . T h e m a x i m u m t h i c k n e s s o f t h i s r h i z o m e

i s s o m e w h a t g r e a t e r t h a n t h a t o f t h e c u l m t h a t g e n e r a t e s .

T h e l a t e r a l b u d s o f t h e r h i z o m e a r e d o m e - s h a p e d a n d g e n e r -

a t e o n l y r h i z o m e s . T h e r h i z o m e i s n a r r o w a t t h e n e c k w h e r e

i t i s a t t a c h e d t o t h e m o t h e r r h i z o m e , a n d t h i c k a n d b r o a d a t

t h e o t h e r e n d w h o s e a p e x p r o t u d e s o u t o f t h e g r o u n d a n d

g r o w s i n t o a n e w c u l m s h o o t w h i c h g e n e r a t e s t h e c u l m . I n

t h e f o l l o w i n g y e a r a n e w r h i z o m e g r o w s , w h i c h i s i s g e n e r a -

t e d b y t h e m e r i s t e m a t i c z o n e o f o n e o f t h e l a t e r a l b u d s o f

t h e m o t h e r b a m b o o .

T h e b u d p r o t r u d e s f o r m i n g f i r s t t h e n e w r h i z o m e n e c k ,

a n d t h e n t h e s h o o t o f t h e r h i z o m e i t s e l f , w h i c h i s c o v e r e d

w i t h s h e a t h s . I n t h e r h i z o m e , t h e s h e a t h s h a v e t h e f u n c t i o n

o f p r o t e c t i n g t h e t e n d e r a p i c a l m e r i s t e m b y f o r m i n g a

l

Part One- Section 1- Bamboo Morphology and Physiology 7

sharp-pointed resistant shield around it to protect it whenthe rhizome is pushed through the soil by the elongation ofthe rhizome internodes during its growing process. Oncedeveloped, (in two or three months, depending on thespecies),the new rhizome turns its distal end or apicalmeristem turns upward and forms the culm shoot.

In the tropics new culm shoots or sprouts begin toappear after the beginning of the rainy season. They pro-trude from the ground as scaly cones covered with sheaths.In warm regions with precipitation at frequent intervalsthroughout the year, the growth is often more or less con-tinuous.Once the culm-shoot is formed, the apical meris-tem stops its function and the intercalary meristems locatedbetween the nodes start the growing process of each one ofthe internodes starting from the lowest one.

In pachymorph species, the distance between the culmsdepends on the length of the rhizome neck and on the posi-tion of the rhizome. When the rhizome neck is short, andthe position of the rhizome is almost vertical, densely caes-pitose clumps of bamboo are formed, as in the genera Bam-busa (Bambusa vulgaris), Dendrocalamus and somespecies of genus Guadua such as "Guadua brasilera", whichis cultivated in Costa Rica .Fig 1.6 B. When the neck of therhizome is long and the rhizome has an almost horizontalposition, as in genera Melocana, Fargesia, and Guaduaangustifolia, the culms grow separatelly and open clumpsare formed. According to Arbelaez (1996) the minimum andmaximum separation beetwen the mother bamboo and thenew culm shoot in a clump of Guadua angustifolia was0.85 to 1.70 meters. In Asia there are species of Melocanna

4-Culms ,

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8 P a r t O n e - S e c t i o n 1 - B a m b o o M o r p h o l o g y a n d P h y s i o l o g y

A c c o r d i n g t o D e o g u n ( 1 9 3 6 ) , t h e r i z h o m e o f D e n d r o -

c a l a m u s s t r i c t u s s h o w s i g n s o f d e t e r i o r a t i o n i n t h e i r f i f t h

y e a r , s o o n e w o u l d n o t e x p e c t n e w c u l m s f r o m r h i z o m e s o f

c u l m s v e y e a r s o r m o r e i n a g e . I

A c c o r d i n g t o B a h a d u r ( 1 9 7 9 ) , t h e r h i z o m e o f D e n d r o -

c a l a m u s h a m i l t o n i i w i t h s l i g h t t r i m m i n g a n d d r e s s i n g i s a n

e x a c t r e p l i c a o f a r h i n o c e r o s h o r n w h i c h f e t c h e s a f a b u l o u s

p r i c e a s a n a p h r o d i s i a c . P e r h a p s o n l y a n e x p e r t c a n i d e n t i -

f i e d t h e i m i t a t i o n r h i n o h o r n f r o m t h e r e a l o n e . T h e n e f a r i -

o u s u s e , h o w e v e r , n e e d s t o b e d i s c o r a g e d . B e c a r e f u l !

C . - M E T A M O R P H R H I Z O M E

K n o w n a l s o a s a m p h i p o d i a l , t h e m e t a m o r p h r h i z o m e

i n c l u d e s t h o s e s p e c i e s w h i c h h a v e t h e c a p a c i t y t o p r o d u c e

b o t h t y p i c a l p a c h y m o r p h a n d t y p i c a l l e p t o m o r p h r h i z o m e s

i n t h e s a m e p l a n t . I n A m e r i c a t h i s i s a c h a r a c t e r i s t i c o f

C h u s q u e a f e n d l e r i ( F i g . 1 . 7 ) a n d o t h e r s s p e c i e s o f t h i s A m e -

r i c a n g e n u s . I n A s i a , a c c o r d i n g t o T i a n ( 1 9 8 9 ) , B a s h a n i a

f a r g e s i i , f r o m C h i n a , p r o d u c e s s h o o t s i n s p r i n g , s u m m e r

a n d a u t u m n , b u t 9 8 . 9 % o f i t s t o t a l s h o o t o u t p u t i s p r o d u c e d

i n s p r i n g .

T h e p e a k p e r i o d o f s h o o t i n g c o m e s w h e n t h e s o i l t e m -

p e r a t u r e a t t h e d e p t h o f 1 0 c m u n d e r g r o u n d r e a c h e s 1 0 t o

1 2 C . T h e f o r m a t i o n a n d e m e r g e n c e o f s h o o t s a r e c l o s e l y

r e l a t e d t o t h e a l t i t u d . W i t h e v e r y r i s e o f 2 0 0 m i n e l e v a t i o n ,

s h o o t i n g b e g i n g s b y 1 3 1 5 d a y s l a t e r .

B o t h t h e r h i z o m e b u d a n d c u l m b a s e l a t e r a l b u d o f t h i s

b a m b o o c o m e u p i n t o s h o o t s d u e t o i t s a m p h i p o d i a l r h i -

z o m e s y s t e m . T h i s r e s u l t s i n t h e m i x e d d i s t r i b u t i o n o f s h o o t

i n t h e s t a n d s . T h e s h o o t s d e v e l o p e d f r o m c u l m b a s e s i d e

b u d s a r e m o r e a n d b e t t e r t h a n t h o s e f r o m r h i z o m e b u d s .

T h e r h i z o m e s y s t e m k e e p s i t s s h o o t i n g p o w e r f o r m o r e t h a n

1 5 y e a r s , w h i l e t h e s i d e b u d s o n l y 5 - 6 y e a r s . T h e b u d s o f 4 -

5 y e a r s o l d r h i z o m e i s m o s t p r o d u c t i v e . T h e s h o o t s a t t a i n e d

t h e i r f u l l h e i g h t i n 9 0 d a y s .

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F i g . 1 . 7 M e t a m o r p h r h i z o m e o f C h u s q u e a f e n d l e r i .

T h i s k i n d o f r h i z o m e c a n p r o d u c e l e p t o m o r p h a n d

p a q u i m o r p h r h i z o m e s i n t h e s a m e p l a n t .

2 . - T H E C U L M

A s m e n t i o n e d a b o v e , i n s p e c i e s w i t h l e p t o m o r p h r h i -

z o m e , t h e c u l m c o n s i s t o f t w o p a r t s : t h e a e r i a l p a r t o r c u l m

p r o p e r a n d t h e s u b t e r r a n e a n p a r t o f t h e c u l m k n o w n a s

c u l m b a s e w i t h i t s r o o t s y s t e m , w h i c h i s c o n e c t e d t o t h e r h i -

z o m e b y t h e c u l m n e c k .

I I n s p e c i e s w i t h p a c h y m o r p h r h i z o m e , t h e c u l m b a s e

w i t h r o o t s d o e s n o t e x i s t a n d t h e a e r i a l p a r t o f t h e c u l m i s

c o n e c t e d d i r e c t l y t o t h e r h i z o m e , w h i c h i n t h i s c a s e c o u l d b e

c o n s i d e r e d a s a c u l m b a s e .

F o r m a t i o n a n d g r o w i n g o f t h e c u l m

b a s e a n d c u l m s h o o t i n s p e c i e s w i t h

l e p t o m o r p h r h i z o m e s

I n s p e c i e s w i t h l e p t o m o r p h r h i z o m e s t h e c u l m i s g e n e r -

a t e d f r o m o n e o f t h e a p i c a l m e r i s t e m s l o c a t e d i n o n e o f t h e

b u d s o f t h e r h i z o m e , w h i c h p r o t r u d e s a n d f o r m s t h e c u l m

n e c k a n d t h e n i t s w e l l s u n t i l i t b e c o m e s a l o n g s h o o t o r

s p r o u t , t h i c k e r t h a n t h e r h i z o m e a n d c o v e r e d w i t h s h e a t h s ,

w h i c h l o o k s l i k e t w o c o n e s j o i n e d b y t h e i r b a s e s ( S e e F i g u r e

1 . 9 ) . C o m p r e s s e d i n t h e s h o r t l o w e r i n v e r t e d c o n e , t h i s s h o o t

i n c l u d e s 1 3 t o 1 6 n o d e s , w h i c h o n c e d e v e l o p e d f o r m t h e

c u l m b a s e . T h e u p p e r c o n e , w h i c h i s l o n g e r , t h a n t h e l o w e r

o n e , c o r r e s p o n d s t o t h e c u l m s h o o t , w h i c h o n c e e l o n g a t e d

b e c o m e s t h e c u l m ( S e e F i g u r e 1 . 8 ) .

A n i n t e r e s t i n g f e a t u r e o f t h i s l o n g s h o o t , i s t h a t a l l t h e

n o d e s o f t h e c u l m b a s e , a n d t h e n o d e s o f t h e c u l m s h o o t t h a t

o n c e e l o n g a t e d w i l l b e c o m e t h e w h o l e c u l m , ( t h e a e r i a l c u l m

a n d t h e c u l m b a s e ) a r e c o m p r e s s e d i n t o i t . I n o t h e r w o r d s ,

m o s t o f t h e n o d e s o f t h e c u l m l i e a t o n e o n t h e t o p o f t h e

o t h e r l i k e a p i l e o f p l a t e s , w i t h t h e p i t h b e t w e n t h e m l a r g e r

a n d t h i c k e r o n t h e b o t t o m , a n d t a p e r i n g o f f t o t h e s m a l l e r a n d

t h i n n e r a t t h e t o p n e a r t h e g r o w i n g p o i n t o r a p i c a l m e r i s t e m .

O n c e t h e a p i c a l m e r i s t e m h a v e f i n i s h e d t h e f o r m a t i o n

o f t h e w h o l e c u l m s h o o t , i t s t o p s i t s f u n c t i o n a n d t h e i n t e r -

c a l a r y m e r i s t e m s l o c a t e d b e t w e e n t h e n o d e s o f t h e c u l m -

b a s e , s t a r t t h e c o n s t r u c t i o n o r e l o n g a t i o n o f t h e i r i n t e r n o d e s

T h e e l o n g a t i o n o f t h e i n t e r n o d e s o f t h e c u l m b a s e i s v e r y

s h o r t b e c a u s e i t s f u n c t i o n i s t o r i s e t h e m a x i m u m d i a m e t e r

o r b a s e o f t h e c u l m s h o o t ( w h i c h s e p a r a t e t h e c u l m p r o p e r

f r o m t h e c u l m b a s e ) u p t o t h e g r o u n d l e v e l . O n c e t h e g r o w i n g

p r o c c e s s o f t h e c u l m b a s e h a s b e e n c o m p l e t e d , a n d t h e b a s e

o f t h e c u l m s h o o t i s a l r e a d y l o c a t e d a t t h e g r o u n d l e v e l ,

s t a r t s i t s g r o w i n g p r o c e s s .

G r o w i n g p r o c e s s o f t h e c u l m s h o o t

i n s p e c i e s w i t h l e p t o m o r p h a n d p a c h y -

m o r p h r h y z o m e s .

T h e g r o w i n g p r o c e s s o f t h e c u l m s h o o t o r a e r i a l c u l m , i s

t h e s a m e f o r s p e c i e s w i t h l e p t o m o r p h a n d p a c h y m o r p h r h i -

z o m e s , b u t i s d i f f e r e n t o f t h a t o f t r e e t r u n k s . U n l i k e t r e e s t h e

b a m b o o c u l m d o e s n o t g r o w i n d i a m e t e r a n d t h e g r o w i n g

p r o c e s s o f t h e c u l m i s c a r r i e d o u t b y t w o k i n d s o f m e r i s t e m s

k n o w n a s a p i c a l m e r i s t e m a n d i n t e r c a l a r m e r i s t e m s . T h e

a p i c a l m e r i s t e m i s i n c h a r g e o f t h e f o r m a t i o n o f t h e r h i z o m e

s h o o t , c u l m b a s e a n d c u l m s h o o t , a n d b r a n c h s h o o t s . T h e

i n t e r c a l a r m e r i s t e m s l o c a t e d b e t w e e n t h e n o d e s , a r e i n

c h a r g e o f t h e g r o w i n g o f e a c h o f t h e s e p a r t s w h i c h h a v e b e e n

f o r m e d b y t h e a p i c a l m e r i s t e m s , i n c l u d i n g t h e g r o w i n g

Part One- Section 1- Bamboo Morphology and Physiology

Fig.1.8 FORMATION AND GROVVTNG PROCESS OF THE CULM BASE AND CULMSHOOT IN LEPTOMORPH BAMBOOS

Culm shoot -

Rhizome

-

of the walls of each one of the intern-odes of the culm shoot starting fromthe lowest.

Once the formation process of theculm shoot is completed by the apicalmeristem, it stops its function and thegrowing process or growing of theinternodes of the culm shoot conti-nues. Thus each internode has its grow-ing zone or intercalary meristem,between the nodes and consequentlyeach internode grows separately, start-ing from the basal internode The, culmelongates joint by joint until it reachesits maximum height. The culm shootceases growing only when the lastinternodal growth is completed. It canbe confirmed that the growth of eachinternode is completed when the culmsheaths peel off slightly from their baseor sheath scar.

Sprout emergence of tropical bam-boos is commonly at the onset of therainy season and in temperate bamboosit is in spring. Nevertheless, it can varyaccording to the species, the vigour ofthe mother bamboo, and the environ-mental condition of the site Even in onebamboo grove, however there are varia-tions of 50 to 60 days between earlyand late sprouting.

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T h e g r o w i n g p e r i o d i s l o n g e r f o r e a r l y s p r o u t s a n d

s h o r t e r f o r t h e l a t e s p r o u t s . E a r l y s p r o u t s t e n d t o d e v e l o p e

i n t o l a r g e r c u l m s w i t h s u p e r i o r q u a l i t y , w h i l e l a t e s p r o u t s

d e v e l o p i n t o s m a l l e r c u l m s w i t h i n f e r i o r q u a l i t y . W h e n t h e

p r e c i p i t a t i o n i s l o w t h e n u m b e r o f s p r o u t s d e c r e a s e s . ( U e d a

1 9 6 0 , L u e t a l 1 9 8 9 ) .

I n N o r t h e r n I n d i a B a m b u s a a r u n d i n a c e a a n d D e n d r o -

c a l a u s s t r i c t u s s e n d u p t h e i r n e w c u l m s h o o t s i n J u n e o r

J u l y w h e n t h e s o u t h w e s t m o n s o o n r a i n s b e g i n s , b u t i n

S o u t h I n d i a t h e n e w c u l m s s h o o t s a p p e a r i n S e p t e m b e r o r

O c t o b e r w i t h t h e n o r t h e a s t m o n s o o n r a i n s .

A c c o r d i n g t o K u r z ( 1 8 7 5 ) i n s o m e p r o v i n c e s o f I n d i a

t h e r e i s t h e b e l i e f t h a t a t h u n d e r s t o r m i s n e c e s a r y b e f o r e t h e

b a m b o o c a n s h o o t . P e r h a p s t h i s i s d u e t o t h e c o i n c i d e n c e o f

t h e r a i n y s e a s o n a n d s t o r m s . T h i s i s c o n f i r m e d b y W a n g

( 1 9 9 3 ) w h o s a y s t h a t t h e C h i n e s e s p e c i e P h y l l o s t a c h y s p r a e -

c o x , s p r o u t s w h e n t h u n d e r o c c u r s i n e a r l y s p r i n g s o i t i s

c a l l e d P h . p r a e c o x , w h i c h m e a n s " t h u n d e r s h o o t " .

W h e n t h e c u l m s h o o t o f g i a n t b a m b o o s a p p e a r s o n t h e

g r o u n d c o v e r e d w i t h s h e a t h s a n d i t h a s a h e i g h t o f a b o u t 3 0

t o 4 0 e m s , t h e c u l m b a s e h a s r e a c h e d i t s m a x i m u m d i a m e t e r ,

a n d t h e r e a f t e r t h i s w i l l n e v e r i n c r e a s e . T h e c u l m s h o o t s t a r t s

g r o w i n g b y e l o n g a t i o n o f t h e i n t e r n o d e s , s t a r t i n g a t t h e l o w e s t

i n t e r n o d e . A t r s t t h e y g r o w s l o w l y , t h e n v e r y f a s t a n d n a l l y

v e r y s l o w l y a g a i n u n t i l t h e y c o m p l e t e t h e i r f u l l g r o w t h . I n

s p e c i e s o f p a c h y m o r p h t y p e t h e w h o l e g r o w t h t a k e s a b o u t 8 0

d a y s , a n d i n g i a n t s p e c i e s s u c h a s G u a d u a a n g u s t 1 f o l i a . u p t o

1 8 0 d a y s . I n s p e c i e s o f t h e l e p t o m o r p h t y p e , i t t a k e s 3 0 d a y s

f o r t h e s m a l l s p e c i e s a n d u p t o 1 3 0 d a y s ( 4 - 5 m o n t h s ) f o r t h e

g i a n t s p e c i e s T h e n i t d e v e l o p s t h e b r a n c h e s a n d l e a v e s a n d

b e c o m e s a m o t h e r b a m b o o b e f o r e c o m p l e t i n g t h e r s t y e a r .

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I f t h e a p e x o f t h e g r o w i n g c u l m s h o o t i s c u t o f f , t h e

e l o n g a t i o n o f t h e r e m a i n i n g p o r t i o n d o e s n o t s t o p . T h i s

s h o w s t h a t w h e n t h e i n t e r c a l a r m e r i s t e m s a r e a c t i v a t e d , t h e

a p i c a l m e r i s t e m i s d e a c t i v a t e d . O n c e t h e f u l l g r o w t h o f t h e

l a s t i n t e r n o d e i s c o m p l e t e d t h e h e i g h t o f t h e c u l m d o n o t

e n c r e a s e a n y m o r e . A t t h i s p o i n t , t h e g r o w i n g p r o c e s s o f t h e

l a t e r a l b r a n c h e s o f t h e c u l m w h i c h a r e g e n e r a t e d f r o m t h e

m e r i s t e m a t i c z o n e l o c a t e d i n t h e c u l m b u d s ( a x i l l a r y m e r i s -

t e m s ) s t a r t s .

T h e b r a n c h s h o o t e m e r g e f r o m t h e b u d s l o c a t e d o n

a l t e r n a t e s i d e s o f t h e c u l m j u s t a b o v e t h e s h e a t h s c a r a t s u c -

c e s s i v e n o d e s , a n d t h e y e l o n g a t e i n t h e s a m e w a y a s t h e

c u l m . O n c e t h e f u l l g r o w t h o f t h e l a s t i n t e r n o d e i s c o m p l e -

t e d , d e v e l o p m e n t o f t h e l e a v e s s t a r t s .

D a i l y g r o w t h o f t h e c u l m - s h o o t

A c c o r d i n g t o s e v e r a l s t u d i e s , t h e d a i l y g r o w t h c o r r e l a t e s

p o s i t i v e l y w i t h t e m p e r a t u r e a n d n e g a t i v e l y w i t h h u m i d i t y .

I n m o s t o f s p e c i e s o f l e p t o m o r p h t y p e s u c h a s t h o s e o f

g e n u s P h y l l o s t a c h y s w h i c h s p r o u t s i n t h e s p r i n g i n J a p a n ,

t h e e l o n g a t i o n o f t h e c u l m s p r o u t o c c u r s m o r e d u r i n g t h e

d a y t h a n d u r i n g t h e n i g h t . H o w e v e r , t h e p a c h y m o r p h b a m -

b o o s i n t r o p i c a l r e g i o n s a r e t h e o p p o s i t e . F o r e x a m p l e , D e n -

d r o c a l a m u s s t r i c t u s g r o w s t w i c e a s m u c h d u r i n g t h e n i g h t

a s d u r i n g t h e d a y , a n d t h e r e a r e s p e c i e s s u c h a s B a m b u s a

o l d h a m i w h i c h g r o w s t h r e e t i m e s m o r e d u r i n g t h e n i g h t

t h a n d u r i n g t h e d a y .

J i n ( 1 9 8 9 ) r e p o r t e d t h a t i n t h e s o u t h e r n r e g i o n o f j i a n g ,

C h i n a , t h e t o t a l g r o w t h o f B a m b u s a o l d h a m i l a s t e d 1 7 1 d a y s

Part One- Section 1- Bamboo Morphology and Physiology 11

(65 days for underground growth and 106 days for above-ground growth). The total growth in this time (or accu-mulative growth) was 712 cm and the mean daily growthwas 6.71 cm.

According to the updated records mentioned by Ueda(1960, 1981) the greatest growth rates of a bamboo culmshoot per day (24 hours) are the following:

a) 91.3 cms by Bambusa arundinacea observed atKew Garden in England in 1855.

b) 88 cm by Phyllostachys edulis (Ph. pubescens)which K. Shibata observed at Koishikawa Botanical Gardenin Tokio in 1898.

c) 119 cm by Mosochiku, Phyllostachys edulis (diame-ter of 16 cm), observed on May 24, 1956 by Koichiro Uedain Nagaoka, Kyoto Prefecture, Japan.

d) 121 cm by Madake, Phyllostachys reticulata (12 cmin diameter) which is the maximum record of daily elonga-tion (24 hours). It was measured by Koichiro Ueda on theoutskirts of Kioto on June 23, 1955.

In Colombia, the maximum elongation observed inGuadua angustifolia has been 30 cm in 24 hours. I haveobserved only elongations of 11 to 15 cm. (Hidalgo 1978).

CHARACTERISTICS OF THE CULMOnce the culm shoot has nished the elongation of the

last internode it becomes a culm. The culm or stem of bam-boos consists of a regular alternation of nodes and inter-nodes, generally hollow, which together with the hard, thickand cylindrical wall gives the culm great mechanical strength.

The nodes are important centers of morphogeneticactivity and intercalary growth. Roots and branches emergeonly at the nodes and in many species they may be swollen.The culm node consists of sheath scar, the supra-nodalridge, the diaphragm and the intranode. In most speciesculm buds emerge from the midle part upwards on alter-nate sides of the culm just above the sheath scar at succes-sive nodes, but in Guadua angustifolia culm buds emergefrom the lower to the top part of the culm

According to Yulong & Liese (1996) The intranode(between the nodal ridge and the sheath scar) can vary inlength between 3-10 mm. The nodadl region of bamboosrevealed that the anatomical structures are basically simi-lar between pachymorph and leptomorph species. Most ofthe bamboos have distinct nodes, which differ in shapeamong the species. Some pachymorph species developdense air roots around the nodal ridge at the bottom part ofthe culm. The form of the diaphragm can vary along oneculm. It may be plane or its central part formed upwards ordownwards or it may even be folded. Furthermore, thenodal structure is important for the liquid movement duringdrying and preservation as well as for the physical andmechanical properties of the culm.

The internodes are delimited by a sheath scar,which marks in the exterior surface the insertion of the culmsheath or culm leaf to the culm. The internodes are shorteron the bottom and top of the culm and are generally longerat the middle part of the culm. For example in Guaduaangustifolia the average lengths are : 12-24 cm. (bottom),40 cm. (middle) and 30 cm (top). There are species such asPleioblastus longinternodios in which the length of theinternodes varies between 70 -94 cms. (Yang 1989). In somespecies the internodes of the base are very long as in Den-

Fig. 1.12 - Nodal and internodal morphology

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A. Nodal Morphology (Liese, 1998)

1 9 2

3'=._ 1.-Guadua angustifolia

5 2.-Guadua cebollaii! awn:-alumna rum-15

I 3. - Dendrocalamus8 - strictus1

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i 4.-Guadua amplexifolia

B. Morphology of internodal bamboo sections

drochloa distans (tamyinwa) of southern Burma, whichmay reach a length of 1.80 meters.

Culms of the same clump and with the same high do nothave the same number of internodes In Colombia, I havefound in two culms of Guadua angustifolia with lengths of17,13 m and 17,36 m., 80 and 76 internodes respectively, andin two culms of 18.40 and 19 meters, 82 and 78 internodesrespectively. This means that there is no relationshipbetween the height of the culm and the number of intern-odes. Some of the internode characteristics are the surfacetexture, the combination and pattern of colors, and the typeof pubescense. The internodes are usually glabrous andsmooth or rough and hairy, becoming glabrous at maturity.When young they are covered with a waxy coating on top. Acharacteristic of many bamboos is the appearance of a whiteexudate on the surface of the culm internodes.

Color of the culms.Not all the bamboo culms are green, some are green

with yellow stripes such as Guadua angustifolia var. bicol-or, which is the most beautiful specie of the Americas. Otherculms are yellow or yellow with green stripes such asPhyllostachys viridis sulphurea and Bambusa vulgarisvar. vittata. It is important to point out that the yellow andgreen colors of this specie were chosen for the Brazilianflag. There are species that are black like Phyllostachys

1 2 P a r t O n e - S e c t i o n 1 - B a m b o o M o r p h o l o g y a n d P h y s i o l o g y

n i g r a , o r b l a c k w i t h y e l l o w s t r i p e s l i k e g u a d u a r a y a d a

a m a r i l l a " ( s t i l l n o t i d e n t i f i e d ) o f C o l o m b i a . A l s o , t h e r e a r e

s m a l l w h i t e a n d r e d s p e c i e s .

A c c o r d i n g t o T a i K a i - c h i ( 2 6 5 - 3 0 6 A . D . ) , i n C h i n a t h e r e

w e r e r e d a n d w h i t e g i a n t s p e c i e s s u c h a s t h e K u e i c h u ( c a s -

s i a b a m b o o ) , w h o s e c u l m s w e r e r e d a n d t h e l a r g e s t o n e h a d

a c i r c u n f e r e n c e o f " t w o c h i " ( 0 . 6 6 m t s ) . T h e w h i t e s p e c i e

k n o w n a s H u a n g c h u ( B u s h y b a m b o o ) , " h a s a s k i n a s w h i t e

a s h o a r f r o s t " . T h e w h i t e s p e c i e w h i c h w a s k n o w n a s P a i l u

c h u , ( w h i t e d e e r b a m b o o ) h a s t h e c h a r a c t e r i s t i c t h a t " i n t h e

h o t s u m m e r s e a s o n , t h e w h i t e s k i n a n d e s h b o t h b e c o m e

r e d " . T h i s v a r i e t y i s f o u n d i n W u l i n g ( C h a n g t e ) .

N a t u r a l c u l m f o r m s

M o s t o f t h e b a m b o o s p e c i e s h a v e r o u n d c u l m s i n s e c -

t i o n b u t t h e r e i s a l s o a s p e c i e w h i c h h a s n a t u r a l s q u a r e

c u l m s , a n d i n a n c i e n t C h i n a t h e r e w e r e a l s o s p e c i e s w i t h t r i -

a n g u l a r c u l m s . I t i s i m p o r t a n t t o p o i n t o u t t h a t t h e c u l m c a n

b e d e f o r m e d a r t i c i a l l y , t r a n s v e r s a l l y a n d l o n g i t u d i n a l l y a s

w e w i l l s e e i n o t h e r c h a p t e r . T h e f o l l o w i n g a r e s o m e o f t h e

u n u s u a l b a m b o o f o r m s :

1 . - T h e n a t u r a l s q u a r e b a m b o o f r o m C h i n a

A c c o r d i n g t o P o r t e r f i e l d ( 1 9 2 5 ) , t h e f i r s t i n f o r m a t i o n

a b o u t t h e s q u a r e b a m b o o T e t r a g o n o c a l a m u s a n g u l a t u s

( P h y l l o s t a c h y s q u a d r a n g u l a r i s ) , a p p e a r e d i n a J a p a n e s e

b o o k " M o K u K i n Y o S i u ( T r e e s a n d s h r u b s w i t h o r n a m e n -

t a l f o l i a g e ) p u b l i s h e d i n K i o t o i n 1 8 2 9 . I t i s s a i d t h a t t h i s

b a m b o o w a s i n t r o d u c e d f r o m C h i n a .

T h e e a r l i e s t r e f e r e n c e s t o s q u a r e b a m b o o ( F a n g c h u ) i n

C h i n e s e l i t e r a t u r e , s a y s t h a t i n t h e y e a r 1 2 8 B . C . i n

S z e c h u a n , w a l k i n g - s t i k s o r " s t a v e s m a d e f r o m s q u a r e b a m -

b o o o f C h i u n g w h i c h n e v e r b r e a k s w e r e u s e d . T h e C h i n e s e

a t r i b u t e t h e s q u a r n e s s o f t h i s p e c u l i a r b a m b o o t o s u p e r n a -

t u r a l p o w e r s .

T h e N i n g p o G a z e t t e e r s a y t h a t a K 0 H s i e n ( f a i r y ) p r e -

p a r e d s o m e e l i x i r o f l i f e w h i c h h e d r a n k a n d a s c e n d e d t h e

m o u n t a i n c a l l e d L i n g f e n g i n T i n g h a i , w h e r e h e p l a n t e d

s o m e c h o p s t i c k s w h i c h g r e w a n d d e v e l o p e d i n t o b a m b o o

w i t h s q u a r e f o r m . ( K u a n g - c h i , 1 6 4 0 )

T h i s b a m b o o g r o w s f r o m 3 " t o 1 0 m e t e r s h i g h . T h e

s q u a r e n e s s i s n o t s o e v i d e n t i n s m a l l e r a n d l e s s m a t u r e c u l m s

a s i t i s i n t h e o l d e r a n d s p e c i a l l y t h e l a r g e r c u l m s . T h i s s p e c i e

i s v a l u e d c h i e y a s a n o r n a m e n t a l g a r d e n p l a n t . I t w a s r s t

d i s c o v e r e d i n t h e f o r m o f s t a v e s b e i n g u s e d b y B u d d h i s t

m o n k s a n d v i l l a g e e l d e r s . T h e s m a l l e r a n d l e s s s q u a r i s h

c u l m s w e r e u s e d f o r o p i u m p i p e s , a n d t h e s t i l l s m a l l e r o n e s

f o r t o b a c c o p i p e s . ( S e e H o w t o m a k e s q u a r e b a m b o o ) .

2 - I n t e r n o d e s w i t h t h e s h a p e o f b o t t l e s

T h e s e a r e f o u n d i n t h e s p e c i e s : D e n d r o c a l a m u s l a t i -

o r u s M u n r o v a r . l a g e n a r i u s L i n . a n d B a r n b u s a v e n t r i c o s a

M c C l u r e . T h i s i s o n e o f t h e m o s t b e a u t i f u l b a m b o o s .

3 . - T r i a n g u l a r . I n t h e b o o k K u a n C h u n F a n g P u ,

p u b l i s h e d i n C h i n a i n 1 7 0 7 , t h e r e i s a d e s c r i p t i o n o f a n

a n c i e n t C h i n e s e b a m b o o k n o w n a s S a n l e n g c h u ( T r i a n -

g u l a r b a m b o o ) . T h e c u l m s o f t h i s v a r i e t y h a d t h r e e a n g l e s .

4 . - F l a t c u l m s . I n t h e s a m e b o o k t h e P i e n c h u ( F l a t

b a m b o o ) i s a l s o m e n t i o n e d . A c c o r d i n g t o S u n p u t h i s v a r i -

e t y w a s g r o w n i n l a r g e n u m b e r s i n t h e K u a n g l u s h a n , a

m o u n t a i n b e t w e e n H s i n g - t z u a n d C h i u c h i a n g d i s t i r i c t s i n

K i a n g s i ) . T h e c u l m a n d t h e n o d e s a r e a t a n d v e r y l o n g . I t i s

s a i d t h a t t h i s v a r i e t y h a s b e e n a t t e n e d i n t o t h i s f o r m b y t h e

B u d d i s t G o d . P r o b a b l y t h i s s p e c i e i s t h e s a m e s q u a r e b a m -

b o o o r a s i m i l a r v a r i e t y .

F r e a k c u l m s

T h e r e a r e s e v e r a l b a m b o o s p e c i e s t h a t s u f f e r a t y p i c a l

d e f o r m a t i o n d u r i n g t h e i r g r o w i n g p r o c c e s s . T h e s e d e f o r m a -

t i o n s a r e k n o w n a s f r e e k s a n d m a k e t h e m v a l u a b l e p i e c e s

u s e d f o r d e c o r a t i v e p u r p o s e s . T h e m o s t i m p o r t a n t v a r i a -

t i o n s a r e t h e f o l l o w i n g :

1 ) . - T u r t l e s h e l l b a m b 0 0 . I n C h i n e s e t h i s d e f o r -

m a t i o n i s r e f e r r e d t o a s L o o H a n n b e c a u s e o f i t s s t r a n g e n e s s

a n d r a r i t y a n d i t i s r e g a r d e d a s a v e h i c l e o f d i v i n i t y . I n

J a p a n i t i s k n o w n a s K i k k o - C h i k u . I t i s c o n s i d e r e d b y

b o t a n i s t a s P h y l l o s t a c h y s p u b e s c e n s v a r . h e t e r o c y c l a f o r m

s u b c o n v e x a .

2 ) . - B u d d a ' S f a c e B a m b o o . T h i s i s k n o w n i n

J a p a n a s B u t s u m e n - C h i c u . I t i s a n u n u s u a l f o r m o f P h y l -

l o s t a c h y s p u b e s c e n s . ( P h . e d u l i s ) F o r m a n y p e o p l e t h e f o r m

o f t u r t l e s h e l l a n d b u d d a ' s f a c e b a m b o o s a r e n o t c l e a r l y

d i s t i n g u i s h e d .

I n b o t h c a s e s , t h e i n t e r n o d e s o f t h e l o w e r p a r t o f t h e

c u l m s o f t h i s t w o f o r m s a r e g r e a t l y r e d u c e d i n l e n g t h a n d

a r e a r r a n g e d o b l i q u e l y i n a z i g z a g f a s h i o n . I n B u d d a ' s f a c e

t h e o b l i q u e a r r a n g e m e n t o f t h e n o d e s i s s u c h t h a t e a c h

n o d e a l m o s t f o r m r i g h t a n g l e s w i t h t h e n o d e s i m m e d i a t e l l y

a b o v e a n d b e l o w i t .

3 ) . - S p i r a l b a m b o o o r " a o b a d a k e " . P l e i o b l a s t u s

g r a m i n e u s f . r n o n s t r i s p i r a l i s ( R a s e t s u c h i k u ) . T h i s s p e c i e

h a s r h i z o m e m e t a m o r p h ( a n p h y p o d i a l ) , c u l m s 2 - 4 m e t e r s

h e i g h t a n d a d i a m e t e r s o f 1 - 4 c e n t i m e t e r s . T h e c u l m s t h a t

s h o o t o u t w i t h l e p t o m o r p h b r a n c h i n g a r e n o r m a l t y p e s , a n d

m a n y o f t h e o n e s w i t h p a c h y m o r p h b r a n c h i n g a r e a b n o r -

m a l . T h e l o n g s p i r a l c u l m s a r e u s e d f o r d e c o r a t i o n . ( O k a -

m u r k a 1 9 8 6 ) .

A b n o r m a l i t y o f t h e i n t e r n o d e s

A c c o r d i n g t o Z a n l g a n d M a ( 1 9 9 1 ) , p l a n t s p e c i e s r e p r o -

d u c e d v i a a s e x u a l m e a n s a r e g e n e t i c a l l y s t a b l e , y e t d u e t o

t h e i r h e t e r o z y g o t e n a t u r e , s o m e v a r i a t i o n s a n d m u t a t i o n s

m i g h t o c c u r d u r i n g t h e p r o c e s s o f a s e x u a l r e p r o d u c t i o n ,

u n d e r c e r t a i n s p e c i c c o n d i t i o n s . M e a n w h i l e , d u e t o t h e

e x t e n s i v e e x i s t e n c e o f c h a n g e a b l e g e n e s , s o m e i n d i v i d u a l

b e c o m e t h e c h i m a e r a o f m u t a t e d a n d u n m u t a t e d g e n e s .

I t i s n o t u n u s u a l f o r v e g e t a t i v e p l a n t s a n d t h e i r p r o g e n y

t o p r o d u c e d i v e r s i f i c a t i o n s , v a r i a t i o n s a n d m u t a t i o n s . P h .

e d u l i s v a r . h e t e r o c y c l a w h i c h h a s a v e r y h i g h o r n a m e n t a l

v a l u e i s a s p e c i e r e s u l t i n g f r o m a v i r u s i n f e c t i o n o f n o r m a l

P h . e d u l i s w h i c h c a u s e s a b n o r m a l i t y o f t h e c u l m s i n t e r n o d e .

D u e t o t h e f a c t t h a t t h i s m u t a t i o n o c c u r s o n l y u n d e r

c e r t a i n s p e c i f i c c o n d i t i o n s , r e v e r s e m u t a t i o n m i g h t o c c u r

w h e n t h e s p e c i f i c c o n d i t i o n s d i s a p p e a r . O t h e r e x a m p l e s

i n c l u d e B a m b u s a v u l g a r i s c u . w a m i n , B a m b u s a v e n t r i c o s a

c u . n a n a , a n d P h y l l o s t a c h y s a u r e a w h i c h h a v e s w o l l e n a n d

i n c l i n e d i n t e r n o d e a t t h e b a s e o f t h e c u l m .

Part One- Section 1- Bamboo Morphology and Physiology 13

Fig.1.13 NATURAL SHAPES OF BAMBOO CULMS

A _ B c

99F:

-Natural square bamboo (Tetragonocalamus angulatus)- Dendrocalamus latitlorus Munro var. lagenarius Lin.

Bambusa ventricosa

Fig. 1.14 FREAK CULMS

A B CA The turtle shell ( Phyllostachys pubescens var. hetero- C-Spiral bamboo ( Pieioblastus gramineus f. monstrispi-cycla form subconvexa). ralis ) (Rasetsu).B -Buddas face bamboo (Phyllostachys pubescens).

|

ii

i

1E

i

,

1 4 P a r t O n e - S e c t i o n 1 - B a m b o o M o r p h o l o g y a n d P h y s i o l o g y

F i g 1 . 1 5 A B N O R M A I . I T Y I N C U L M S O F G u a d u a a n g u s t i f o l i a

O n t h e C o t o v e f a r m b e l o n g i n g t o t h e N a c i o n a l U n i v e r s i t y i n t h e S t a t e o f A n t i o q u i a , n e a r t h e c i t y o f S a n t a F e d e A n t i o -

q u i a , t h e r e i s a s m a l l p l a n t a t i o n ( 0 . 3 3 h e c t a r e s ) o f G u a d u a a n g u s t i f o l i a A c c o r d i n g t o A n a c i / i a A r b e l a e z , i n t h e c e n t r a l

p a r t o f t h e i n t e r n o d e s , m o s t o f t h e c u l m s h a v e l i k e a w o o d e n r o d w i t h a n a v e r a g e d i a m e t e r o f a b o u t 5 m m w h i c h c o n -

n e c t s s e v e r a l d i a p h r a g m s a l o n g t h e c u l m s l e n g t h , b u t i n s o m e i n t e r n o d e s t h e r o d d i s a p p e a r .

H o w e v e r , t h e s e c h a r a c t e r i s t i c s a r e s o m e t i m e s t a b l e

a n d s o m e t i m e u n s t a b l e i n t h e i r p r o g e n i e s . T h e s e g r e g a t i o n

o f t h e p r o g e n i e s o f t e n m a k e s p r o d u c e r s v e r y u n c e r t a i n . T h i s

i s c a u s e d b y t h e i n s t a b i l i t y o f t h e g e n e t i c f a c t o r s , o r t h e

e n v i r o n m e n t a l c o n d i t i o n s w h i c h p l a y a l e a d i n g r o l e i n t h e

o c c u r e n c e o f s u c h d i v e r s i f i c a t i o n s w h i c h n e e d c o n v i n c i n g

t e s t d a t a t o b e p r o v e n .

T h e r e f o r e , n o o n e i s a b l e t o t a k e t h e i n i c i a t i v e i n p r o -

d u c t i o n . I n a d d i t i o n t h e r e a r e c h a n g e a b l e g e n e s i n t h e v e g e -

t a t i v e p l a n t s w h i c h p r o d u c e c h i m e r a s . T h i s c a n b e s e e n i n

t h e c h i m e r a s o f v a r i o u s c o l o r s o n t h e c u l m s , b r a n c h e s a n d

l e a v e s o f m a n y b a m b o o s p e c i e s s u c h a s P h y l l o s t a c h y s

b a m b u s o i d e s f . t a n a k a e M a k i n o , P h . g l a u c a f . y o u z h u L u

w h i c h h a v e s p o t s o r s t r i p s o f v a r i o u s c o l o r s o n t h e c u l m s , o r

o n t h e c u l m s , b r a n c h e s a n d l e a v e s . H u m a n b e i n g s o f t e n

m a k e u s e o f t h e s e v a r i a t i o n s t o b r e e d n e w b a m b o o s p e c i e s

o f v a r i o u s o r n a m e n t a l v a l u e s .

M a n y s p e c i e s o f t h e g e n e r a P h y l l o s t a c h y s , B a m b u s a ,

e t c . h a v e m a n y v a r i e t i e s o r m u t a n s w h i c h p o s s e s s s t r i p s o f

v a r i o u s c o l o r s . F o r e x a m p l e t h e n o r m a l c o l o r o f t h e c u l m s ,

b r a n c h e s a n d l e a v e s o f P h . e d u l i s a r e g r e e n . P h . e d u l i s f

l u t e o s u l c a t a i s a s p e c i e o f g e n e m u t a t i o n w h i c h h a s r e g u l a r

y e l l o w s t r i p s o n t h e c o n c a v e s i t e o f t h e i n t e r n o d e s . P h .

e d u l i s f . n a b e s h i m a n a h a s y e l l o w a n d g r e e n s t r i p s o n

c u l m s , b r a n c h e s a n d l e a v e s . P h . a u r e o s u l c a t a , P h . b a m b u -

s o i d e s , P h . p r a e c o x , P h . v i v a x , B a m b u s a g l a u c e s c e n s , B .

v u l g a r i s , a l l h a v e s i m i l a r v a r i a t i o n s .

M A T H E M A T I C A L R E L A T I O N -

S H I P S O F T H E C U L M

B a m b o o i s t h e o n l y p l a n t w h i c h h a v e a m a t h e m a t i c a l

r e l a t i o n s h i p b e t w e e n t h e c i r c u n f e r e n c e a n d t h e i n t e r n o d e

a n d t h e c i r c u m f e r e n c e a n d t h e l e n g t h o f t h e c u l m , w h i c h

p e r m i t s n o t o n l y t h e i d e n t i f i c a t i o n o f t h e s p e c i e s b u t a l s o

t h e d e t e r m i n a t i o n o f t h e c u l m h e i g h t .

1 . - I d e n t i c a t i o n o f b a m b o o s p e c i e s

b y m a t h e m a t i c a l r e l a t i o n s h i p .

D i c k a s o n ( 1 9 4 1 ) d e v e l o p e d a n i n t e r e s t i n g t h e o r y r e l a t -

e d t o t h e i d e n t i f i c a t i o n o f t h e s p e c i e s u s i n g a s d i a g n o s i s t h e

m a t h e m a t i c a l r e l a t i o n s h i p w h i c h e x i s t s b e t w e e n t h e c i r c u n -

f e r e n c e o r c u l m g i r t h " C " t a k e n a t a h e i g h t o f 1 . 3 5 m e t e r s

( 4 . 5 f e e t ) a b o v e t h e g r o u n d , a n d t h e l e n g t h o f t h e i n t e r n o d e

" L " l o c a t e d a t t h i s e l e v a t i o n ( 1 . 3 5 m e t e r s ) .

H e g i v e s t h e f o l l o w i n g r e l a t i o n s C / L t a k e n i n s o m e

s p e c i e s : B a m b u s a w a r n i n g B r . 1 . 4 ; D e n d r o c a l a u s g i g a n -

t e u s M u n . 1 . 2 ; T h y r s o s t a c h y s s i a m e n s i s G a m . 0 . 9 4 ; B a m -

b u s a v u l g a r i s S c h r a d . 0 . 9 2 ; B a m b u s a v u l g a r i s v a r . s t r i a t a

R i v . 0 . 9 0 ; D e n d r o c a l a m u s m e m b r a n a c e u s M u n . 0 . 7 0 .

M e l o c c a n n a b a m b u s o i d e s T r i n . 0 . 4 4 ; C e p h a l o s t a c h y u m

p e r g r a c i l e M u n . . 0 . 3 5 ; M e l o c a n n a h u r n i l i s K u r z 0 . 1 7 .

H e s u g e s t e d t h i s h e i g h t ( 1 . 3 5 m t s ) a b o v e t h e g r o u n d

a s a m a t t e r o f c o n v e n i e n c e : i t i s e a s y t o r e a c h , i t i s u s u a l l y

n o t c o v e r e d b y p e r s i s t e n t o v e r l a p p i n g s h e a t h e s , d o e s n o t

r e q u i r e t h e c u t t i n g o f t h e c u l m .

Part One- Section 1- Bamboo Morphology and Physiology 15

2.-Approximate mathematical de-termination of the culms height

There is a mathematical relation between the culmheight and the girth or circumference of the culm taken atthe eye level, as is traditional in Japan; but it can be takenat the breast level (the difference between the girth at breastlevel and that at eye level is very small).

Most buyers of bamboo culms in Japan, multiply theculm girth at eye level by a constant = 60 to obtain the culmlength, or they multiply the diameter by "pi" (3.1416) andthen by 60. This procedure is applicable to the culms over 7cm. in diameter. In Japan thi method has used as means tojudging the quality of the culms. If the culm is longer or thesame length as that determinated mathematically, the culmis of very hihgt quality. If it is smaller then it is very lowquality and consequently has a lower cost.

The constant varies according to the species. In Colom-bia I cut about 60 culms of Guadua angustifolia for myexperiments. With Prof. Jorge Escobar we took all themeasurements of each culm in order to nd the "constant"that we had to use in order to calculate approximately theheight of a culm in this specie, and the result was 58.2.

For example, if the girth at breast level is 0.40 mts. theaproximately height of the culm will be 0.40 x 58.2 = 23.28m. Using the diameter instead of the girth, the height willbe: diameter x 3.1416 x 58.2.

Approximate dimensions of somegiant bamboos of genus Guadua at thetime of Columbus arrival in the Ame-ricas in 1502 .

Using on the above mathematical relation which allowsus to determine approximately the height of the culms ofgiant species of Genus Guadua using the girth or circumfer-ence of the culm, it is possible to calculate approximatelythe-height of the giant culms of genus Guadua, which therewere in the Americas at the arrival of Colubus, ve hundredyears ago. For this purpose we can take in account the infor-mation which exist about the dimensions of their circumfer-ences at that time.

For example: in 1526, Oviedo, a Spanish historian,wrote in ancient Spanish, about the circunferences of bam-boo species in Ecuador: "But among all types of bamboo,there are some that are extremely large and the intern-odes are as big as a heavy mans thigh, and they are threepalms or more in circumference (Patio 1975). - Thepalm is an old Spanish measurement = 23 cm=9 inches. Ifthe largest bamboo in Ecuador (Guadua angustifolia) hada circumference of 3 palms or more. this means that theminimun circumference was of 3 palms = 0.69 m, whichcorresponds to a diameter of 0.22 m.Then the minimumhigh of this bamboo was 0.69 x 58.2 (constant) =40 meters.If we compare these dimensions with those that this speciehas today: 0.10- 0.12 m in diameter by 18 to 21 m hight; thismeans that the dimensions of Guadua angustifolia havebeen reduced by 50% in almost 500 years.

In discussing to the dimensions of the internodes,Simon (1637) points out that "The largest internodes havethe capacity of one arroba of water The arroba is a Span-ish liquid measure which is equal to 25 pounds: 11,500 gr.=

11,500 cm3 of water capacity.If the exterior diameter was 22 cms. the interior was

about 18 cms.This means that the length of the internodewas about 48-50 cms. Probably was taken of the center ofthe culm where are found the largest internodes.

Marco Polo said than on his trip to India in 1290, hesaw very large bamboos with a diameter of 45 centimeterswhich were used as boats by the natives for going from oneside to the other of the largest rivers.

Without doubt he was refering to the specie Dendro-calamus giganteus, the largest bamboo native to India.According to Prosea (1995), this bamboo today is 18-25 cmsin diameter and 26- 30 meters height. This means that thisspecie also has reduced its dimensions by 50% in 710 years.

According to the above, the diameter and height of thebamboo culms in Asia and the Americas have been reducedlittle by little, probably due to continuous climatic changesin the world and to the destruction of the bamboo forests inthe last 200 years.

The huge dimensions of giantbamboos in ancient China

Up until now I have been very sceptical about theinformation I had found in several ancient books fromChina about the extraordinary and incredible dimensionswhich some bamboos in that country had 3000 years ago.But keeping in mind the reduction of the dimensions ofthe culms of most of the giant species in the world, I thinkthat this informations could be valid.

For example, according tothe book Sheng iching(Records related to the marvelous things) written by TungFang, who lived in the 2nd century B.C; in the southernwildernesses in China there was a bamboo known as Ti chu(Weeping bamboo) which grew to a height of several hun-dreds of chan (one chan: 10 feet=3,33 m; one Chinese footor chi = 0.333 m). The circumferece of this bamboo was 3chan and 6 chi; and the culm thickness was eight or ninetsun. (tsun =one inch). Its shoots are delicious and ifthey are eaten they will cure ulcers

If the dimensions have not changed since ancienttimes and if there is no mistake in the dimensions given byHagerty (1919) in his translation of this book, this bamboohad a circunference of three chan antd six chi=36 Chine-se feets =12 m, which corresponds to a diameter of 3.80 m.,and the thickness of the wall would be about 20-22 cm.

The book Chu Pu written by Tai Kai Chi (265-306) saysthat "in Yuangchiu a bamboo known as Tin Chun Chugrows. This bamboo is so large that a boat can be madefrom each one of their internodes , this is conrmed alsoby Kuo Po of Erh ya.

The Treatrise on Agriculture (Nung Cheng ChuanShu) compiled by Hsu Kuang-chi and originally publishedin 1640 says that the Pei chu (Giant bamboo) is found inthe southwestern part of the Han-shan district in Anhuipro-ince. According to the Tzu yuang, this variety mea-sures one-hundred chan (= 333 m in height), has a cir-cumference of three chan and ve or six chi (about 35 chi= 11.65 m = 3,70 m in diameter. The thickness of the culmwall is eight or nine tsun. Its lumber can be used for build-ing boats. Its seeds have a good avor and, if they are eatenthey will cure ulcers or sores.

1 6 P a r t O n e - S e c t i o n 1 - B a m b o o M o r p h o l o g y a n d P h y s i o l o g y

T h e s a m e T r e a t r i s e o n A g r i c u l t u r e ( 1 6 4 0 ) , s a y s t h a t t h e

L u n g k u n g c h u ( D r a g o n d u k e b a m b o o ) i s a l a r g e v a r i e t y

t h a t h a s a d i a m e t e r o f s e v e n c h i ( 1 c h i = 0 . 3 3 3 m x 7 =

2 . 3 3 m i n d i a m e t e r ) a n d b e t w e e n e a c h j o i n t ( i n t e r n o d e )

t h e r e i s a s p a c e o f o n e c h a n ( t e n C h i n e s e f e e t ) a n d t w o

c h i ( 1 2 c h i = o . 3 3 3 x 1 2 = 4 m ) . T h e l e a v e s o f t h i s v a r i e t y

a r e l i k e t h o s e o f t h e C h i a o ( b a n a n a ) w h i c h g r o w s i n t h e L o

f o u s h a n ( r a n g e o f m o u n t a i n s i n t h e K u a n g t u n g p r o v i n c e ) .

I n t h e b o o k T u S h u C h i C h e n g t h e r e i s i n f o r m a t i o n

a b o u t t w o v a r i e t i e s f r o m t h e F a n y u h s i e n d i s t r i c t w h i c h

h a v e t h e s a m e c h a r a c t e r i s t i c s a s t h e a b o v e s p e c i e s , b u t w i t h

d i f f e r e n t n a m e s , t h e D r a g o n d u k e b a m b o o i s k n o w n a s

K i n g o f Y u e h b a m b o o ( Y u e h w a n g c h u ) w h i c h h a s a c u l m

s o t h i c k t h a t i t r e q u i r e s a b o u t f o u r o r v e p e o p l e w i t h

j o i n e d h a n d s t o e n c i r c l e i t . T h i s v a r i e t y o f b a m b o o p r o d u c e s

s h o o t s i n t h e f o u r t h a n d f t h m o n t h s , t h e v o l u m e o f w h i c h

a r e t h e s i z e o f a t o u ( t o u = 1 o 0 l i t e r s ) .

T h e o p p o s i t e i n d i m e n s i o n s i s t h e L u n g s u n c h u

( D r a g o n s g r a n d s o n b a m b o o ) w h i c h g r o w s i n C h e n - c h o u

( l o c a l i t y i n H u n a n ) u s s u a l y i n t h e v a l l e y s a n d r a v i n e s . T h e y

a r e a s n e a n d s l e n d e r a s a n e e d l e w i t h a h e i g h t t h a t d o e s

n o t e x c e e d 3 0 c m .

T h e i n u e n c e o f t h e c u l m a g e

T h e i n u e n c e o f t h e c u l m s a g e o n i t s e c o n o m i c v a l u e i s

p e r h a p s g r e a t e r i n b a m b o o s t h a n i n a n y o t h e r t i m b e r . T h e

a g e o f t h e c u l m w h e n c u t h a s a n i m p o r t a n t b e a r i n g o n i t s

u s e i n t h e m a n u f a c t u r e o f i n d u s t r i a l p r o d u c t s a n d a s m a t e -

r i a l f o r c o n s t r u c t i o n .

A g e i s a v e r y i m p o r t a n t f a c t o r f o r t h e d e v e l o p m e n t o f

c e r t a i n s t r e n g t h p r o p e r t i e s . I t i s t h e g e n e r a l a s s u m p t i o n

t h a t b a m b o o c u l m s m a t u r e i n a b o u t t h r e e y e a r s a n d h a v e

t h e n r e a c h e d t h e i r m a x i m u m s t r e n g t h .

A c c o r d i n g t o L i e s e ( 1 9 8 5 ) , i n v e s t i g a t i o n s w i t h

D e n d r o c a l a m u s s t r i c t u s h a v e s h o w n t h a t i n t h e g r e e n c o n -

d i t i o n , o l d e r b a m b o o c u l m s h a v e h i g h e r s t r e n g t h p r o p e r t i e s

t h a n y o u n g e r o n e s . A c c o r d i n g t o m y e x p e r i e n c e s w i t h

G u a d u a a n g u s t i f o l i a t h e a g e o f t h e c u l m i n c r e a s e s o n l y c e r -

t a i n s t r e n g t h p r o p e r t i e s , s u c h a s c o m p r e s s i o n s t r e n g t h a n d

s h e a r i n g s t r e s s , b u t i n t h e c a s e o f t e n s i l s t r e n g t h t h e r e i s

l i t t l e d i f f e r e n c e b e t w e e n y o u n g b a m b o o s o n e o r t w o y e a r s

w i t h o l d e r o n e s . 1

I n t h e e x p e r i m e n t s I d i d w i t h b a m b o o s t r i p s t a k e n f r o m

t h e e x t e r n a l p a r t o f t h e c u l m w a l l , t h e m a x i m u m t e n s i l

s t r e n g t h w a s 3 , 2 1 7 k g / c m 2 f o r c u l m s 3 . 5 y e a r s o l d , a n d

3 , 2 0 6 k g / c m 2 f o r c u l m s o n e y e a r a n d a h a l f o l d . P r o b a b l y

t h i s i s t h e r e a s o n w h y t h e C h i n e s e u s e d b a m b o o s t r i p s o n e

y e a r o l d o r l e s s f o r t h e m a n u f a c t u r e o f b a m b o o c a b l e s

w h i c h t h e y u s e f o r t h e c o n s t r u c t i o n o f s u s p e n s i o n b r i d g e s

w i t h m o r e t h a n o n e h u n d r e d m e t e r s o f s p a n .

T h e d u r a b i l i t y o f a n i n d i v i d u a l c u l m v a r i e s a c c o r d i n g

t o t h e s p e c i e a n d t h e m a x i m u m i s a b o u t 1 2 y e a r s a t t h e e n d

o f w h i c h t h e c u l m d i e s a n d b e c o m e s d r y a n d w h i t e .

A c c o r d i n g t o m y o b s e r v a t i o n s t h e s t r e n g t h o f G u a d u a

a n g u s t i f o l i a s t a r t s d e c r e a s i n g i n t h e g r o v e w h e n i t i s a b o u t

7 t o 8 y e a s o l d , c o n s e q u e n t l y i t i s r e c o m m e n d e d t h a t c u l m s

u p t o 6 y e a r s o l d b e u s e d i n a l l t h e e l d s o f c o n s t r u c t i o n .

D e t e r m i n a t i o n o f t h e c u l m s a g e

U n l i k e t r e e s , t h e d i a m e t e r a n d h e i g h t o f a b a m b o o

c u l m d o n o t d e t e r m i n e i t s a g e , b u t c e r t a i n c h a n g e s a r e g e n -

e r a l l y r e c o g n i z a b l e a s t h e c u l m b e c o m e s o l d e r . T h e o n l y

m e t h o d f o r d e t e r m i n i n g t h e a g e o f b a m b o o c u l m s a c c u r a t e -

l y , i s m a r k i n g e a c h c u l m s h o o t u s i n g a n a l u m i n u m p l a t e w i t h

a c o l l a r ( n y l o n ) l o c a t e d a r o u n d t h e s h o o t w i t h t h e y e a r a n d

d a t e o f i t s e m e r g e n c e .

I n t h e s e c o n d m e t h o d , " v i s u a l f a c t o r s " t h a t a r e n o t v e r y

e x a c t a r e u s e d . I n t h i s m e t h o d t h e o n e y e a r o l d c u l m s o f

m a n y g i a n t s p e c i e s o f g e n u s G u a d u a a r e e a s i l y r e c o g n i z a b l e

b y t h e e m e r a l g r e e n c o l o r o f t h e c u l m , a n d b e c a u s e t h e y s t i l l

h a v e t h e l o w e r s h e a t h s p a r t i a l l y f i x e d t o t h e c u l m i n i t s l o w e r

p a r t .

T h e t h r e e y e a r s o l d c u l m s c a n b e r e c o g n i z e d b e c a u s e t h e

g r e e n c o l o r i s n o t s o d a r k a n d s m a l l w h i t e d o t s a b o u t 1 - 2 m m

a p p e a r o n t h e s u r f a c e W i t h t h e y e a r s t h e s e i n c r e a s e i n d i a m -

e t e r a n d b e c o m e l a r g e l i c h e n s w h e n t h e c u l m s a r e 5 t o 6

y e a r s o l d . O n e o f t h e b e s t m e t h o d s c o n s i s t s i n c o u n t i n g t h e

l e a f s c a r s i n t h e b r a n c h e s a s i s s h o w n i n t h e F i g . 1 . 1 6

i t A . = O n e y e a r o l d

X l l 7 B . = T w o y e a r s o l d

C . = T h r e e y e a r s o l d

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F i g . 1 . 1 6 D e t e r m i n a t i o n o f t h e c u l m a g e b y n u m b e r o f

l e a v e s s c a r f s i n t h e b r a n c h e s . T h e t o p b r a n c h ( A ) b e l o n g s t o

a b a m b o o o n e y e a r o l d . T h e b r a n c h e s ( B a n d C ) b e l o n g t o

p l a n t s t w o a n d t h r e e y e a r s o l d r e s p e c t i v e l y

\ 1 L e a f s c a r f S 3 9

Part One- Section 1- Bamboo Morphology and Physiology 17

CULM BRANCHESBranchesare generated from the branch bud, or axillary

meristem, which are arranged on alternate sides of theculm. Each internode bears a branch bud (primordium)located just above the culm leaf scar, and may be foundfrom the lowermost node upwards as in Guadua angustifo-lia, or from the midculm node upwards. In some speciesbranches develop while the culm is still growing, in othersthey develop only after the culm reaches its full height.

In some bamboos the branch bud is solitary. Thebranch system is often very characteristic of bamboogenus. If the apex of the culm is damaged, the dormant pri-mary branch axis of each node will grow and elongate toreplace the main culm. The array of branches may developat a single culm node and is called a branch complement. Inbamboos whose culms are branched, it is at the mid-culmnodes than the typical form of the branch complementappears.

The number of branches on each node of the mid-culmsis important taxonomically and it differs with the genus.The branch complement appears most typically at the mid-culm nodes. The lowermost and uppermost branch comple-ment are generally less well developed than those at themid-culm nodes. In bamboo plants there is only one bud ateach node in all genera.

About the branch primordium and its growth . In many

.

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87'"o E F

Fig.1.17 Examples of diverse basal proliferation inthe mid-culm branch complement in several genera:(A) Guadua angustifolia, (B) Phyllostachys dulcis, (C)Shi-bataea kuma-sasa, (D) Arundinaria tecta, (E)Arundinaria simonii, (F) Semiarundinaria fastuosa.McClure (1966).

bamboos the branch primordium ramies precociously inthe prophyll at the early stage. In Sasa, Sasamorpha andmost Sasaella, the primordial branch axis possesses nobasal buds and does not ramify, so the branch remains soli-tary. Phyllostachys typically ramifyes one time and is bina-ry; the primary and secondary branches being unequal,some times a third much smaller one develops between theother two. The primary branch emerges, and remainsstrongly dominant. Subsequent orders of axes that developfrom its basal buds gradually become smaller in size andshorter in length.

THE LEAVESProbably the specie which have the largest leaves is

Neurolepis elata from the Andes which may reach a lengthof 5 meters and a width of 0.50 m. While the smallest arethose of Arthrostylidium capillifolium which are no witherthan 3 mm.

According to Ueda (1987) , bamboo leaves fall off atone year after they appear and are renewd. Old leaveschange into new leaves at the same time. Even the seasonand the way of renewal are different according to thespecies. The season for the species of large size bamboos isspring in general. This special ability for quick renewal isefficient for staying greeen and heightening the efficiencyof photosynthesis.

That is because a bamboo ought to be strong in airpolution. In general the small species also stay green bygradually changing old leaves for new ones as a unit of indi-vidual living bamboos.

In japan leaf change of new bamboos with pachymorphrhizome is nished in July. This is the period for vigorousgrowth of new leaves and is also the period for new rhizomegrowth. There is a close relationship between the leafchange and shoot sprouting. The number of sprouts is lessin leaf changing year, but increases the next year.

Nonaka &Sasaki (1992), observed that in the damage ofbamboo groves in Fukuoka, Japan, by a typhoon in 1991leaves lost color and withered in a few days just after thetyphoon but about ten days later new leaves appeared.

Uses.-Bamboo leaves are used for thatching housesand provide valuable fodder for cattle and for elephants inIndia. According to Kawase & Ujie (1987), in Japan, theleaves of miyakozasa, or tini Sasa, have been used as thefeed for pastured horses since old times. Dried maturebamboo leaves are used for deodorising fish oils. At thesame time, there has been the folklore belief that theextracts from the leaves are effective for the treatment ofsome cancers, and an infusion prepared from them hasbeen drunk.

Since the cancer became a leading cause of mortallityin Japan, the medicinal effect of the extract from Sasa leaf(called Bamfolin ) has been brought to the atention of themedical and farmaceutical elds. A lot of useful results arereported by the tests of animals receiving transplanted can-cer cells. In relatively few clinical tests, Oshima treated 69patients of seriously cancerous diseases with Bamfolin forover six months and reported that some curing effects wererecognized by 10% of all patients, besides a promotion ofappetite and an alleviation of pain. (See Pharmaceuticaluses of leaves).

1 8 P a r t O n e - S e c t i o n 1 - B a m b o o M o r p h o l o g y a n d P h y s i o l o g y

, = , 1 i .

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9 .

S c h i z o s t a c h y u m P h y l l o s t a c h y s D i n o c h l o a

F i g . 1 . 1 8 T y p e s o f c u l m s h e a t h s o r c u l m l e a v e s . 1 . - U p p e r p a r t o f t h e s h e a t h o f B a m b u s a v u i g a r i s s h o w i n g

o n e p a n o f t h e s t r u c t u r e s t h a t a r e i m p o r t a n t i n c l a s s i f i c a t i o n a n d i n i d e n t i f i c a t i o n o f t h e d i f f e r e n t b a m b o o s p e c i e s : ( a )

S h e a t h p r o p e r , ( b ) S h e a t h b l a d e , ( c ) L i g u l e , ( d ) P a i r o f a u r i c l e s . 2 . - U p p e r p a n o f t h e s h e a t h o f G u a d u a a n g u s t i f o l i a i n

w h i c h i t c a n b e s e e n t h a t t h e t r i a n g u l a r b o r d e r i s c o n t i n u o u s a r o u n d t h e s h e a t h . 3 . - S o m e d i f f e r e n t t y p e s o f s h e a t h s .

C u l m s h e a t h ( C u l m l e a f )

A s w a s e x p l a i n e d b e f o r e , t h e g r o w i n g o f t h e b a m b o o

c u l m t a k e s p l a c e i n t h e i n t e r n o d e s . T h e u p p e r p a r t o f e a c h

n o d e c o n s i s t s o f m e r i s t e m a t i c t i s s u e , s u r r o u n d e d b y t h e

c u l m s h e a t h , w h i c h h a s t h e f u n c t i o n o f p r o t e c t i n g , s u p p o r t -

i n g a n d s t i f f e n i n g t h e t e n d e r m e r i s t e m a t i c z o n e o f t h e

i n t e r n o d e s o f t h e c u l m a n d b r a n c h e s d u r i n g t h e i r g r o w i n g

p r o c e s s . I n t h e c u l m s h e a t h t h e r e a r e m a n y v a s c u l a r b u n d l e s

r u n n i n g l o n g i t u d i n a l l y i n a l m o s t p a r a l l e l l i n e s .

A c c o r d i n g t o M c C l u r e ( 1 9 6 6 ) , w h e n C h i n e s e g a r d e n e r s

w i s h t o d w a r f a b a m b o o , t h e y r e m o v e e a c h c u l m s h e a t h p r e -

m a t u r e l y , b e g i n n i n g w i t h t h e l o w e s t , b e f o r e t h e e l o n g a t i o n

t a k i n g p l a c e a b o v e i t s n o d e i s c o m p l e t e d . U p o n t h e r e m o v a l

o f a s h e a t h , t h e e l o n g a t i o n a b o v e i t s n o d e c e a s e s .

T h e c u l m - s h e a t h i s a n i m p o r t a n t s t r u c t u r e f o r b a m b o o

c l a s s i f i c a t i o n . T h e s h a p e , s i z e , t h i c k n e s s a n d h a i r i n e s s o f

b o t h t h e s h e a t h p r o p e r a n d s h e a t h b l a d e a n d t h e e x i s t e n c e

o f a u r i c l e o r o r a l s e t a e a r e i n v a l u a b l e f o r r e c o g n i z i n g t h e

g e n u s o r s p e c i e s . T h e i n s i d e f a c e i s s m o o t h a n d s h i n y ; w h e n

y o u n g t h e o u t s i d e o r t h e b a c k i s u s u a l l y c o v e r e d w i t h i r r i -

t a n t h a i r s w h i c h m a y b e w h i t e , p a l e b r o w n , g o l d e n b r o w n o r

b l a c k . T h e s h e a t h a l s o v a r i e s i n c o l o r a n d m a y b e g r e e n ,

b l u i s h o r p u r p l i s h - g r e e n , o r y e l l o w t i n g e d w i t h o r a n g e .

C u l m s h e a t h s g e n e r a l l y s h e d t h e c u l m u p o n a t t a i n i n g

m a t u r i t y , b u t m a y a l s o b e p e r s i s t e n t ( S a s a , P l e i o b l a s t u s ) ,

l a t e d e s i d u o u s ( I n d o c a l a m u s S e m i a r u n d i n a r i a , F a r g e s i a

e t c . ) o r e a r l y d e c i d u o u s ( B a m b u s a , P h y l l o s t a c h y s , D e n d r o -

c a l a m u s e t c . ) . I n t h e e x t e r n a l f o r m , t h e s h e a t h s o f G u a d u a

a n d C h i m o n o b a m b u s a a r e t r i a n g u l a r , w h i l e t h o s e o f D e n -

d r o c a l a m u s a n d P h y l l o s t a c h y s a r e b e l l - s h a p e d , a n d f o r

S c h i z o s t a c h y u m a n d L i g n a n i a , t h e y a r e l a n c e o l a t e . S h e a t h

l e a v e s o f C h i m o n o b a m b u s a a r e s h o r t a n d s m a l l , a n d a w ] -

s h a p e d . ( K e n g a n d W e n , 1 9 9 1 ) .

U s e s : B a m b o o s h e a t h s p r o v i d e s u b s t i t u t e m a t e r i a l f o