Plant Physiol. 1977 Potter 10 4

8/12/2019 Plant Physiol. 1977 Potter 10 4

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P l a n t P h y s i o l . 1 9 7 7 ) 5 9 , 10-14

L e a f A r e a P a r t i t i o n i n g as an I m p o r t a n t F a c t o r i n GrowthR e c e i v e d f o r p u b l i c a t i o n S e p t e m b e r 1 9 , 1 9 7 5 and i n r e v i s e d form J u l y 1 9 , 1976

J O H N R . P o T r E RS o u t h e r n Weed S c i e n c e L a b o r a t o r y , 2 A g r i c u l t u r a l R e s e a r c h S e r v i c e , U n i t e d S t a t e s Department o f A g r i c u l t u r e ,S t o n e v i l l e , M i s s i s s i p p i 38776J A M E S W. J O N E SC o t t o n P r o d u c t i o n I n s t i t u t e , A g r i c u l t u r a l R e s e a r c h S e r v i c e , U n i t e d S t a t e s Department of A g r i c u l t u r e ,M i s s i s s i p p i S t a t e , M i s s i s s i p p i 39762

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

new l e a f a r e a i s a n i m p o r t a n t c o m p o n e n t o f g r o w t h . A c c o r d i n g l y , a ne x p r e s s i o n w a s d e v e l o p e d t o m e a s u r e l e a f a r e a p a r t i t i o n i n g . U s i n gg r o w t h a n a l y s i s t e c h n i q u e s , r e l a t i v e g r o w t h r a t e s w e r e c o m p a r e d t o n e ta s s i m i l a t i o n r a t e s , p a r t i t i o n i n g o f d a i l y w e i g h t g a i n i n t o new l e a f a r e a ,a n d p a r t i t i o n i n g o f d a i l y w e i g h t g a i n i n t o new l e a f w e i g h t o f n i n e s p e c i e sg r o w n i n g r o w t h c h a m b e r s u n d e r t h r e e t e m p e r a t u r e r e g i m e s . D a y / n i g h tt e m pe ra t u r e s o f 2 1 / 1 0 , 3 2 / 2 1 , a n d 3 8 / 2 7 C c a u s e d l a r g e d i f f e r e n c e s i nr e l a t i v e g r o w t h r a t e s . R e l a t i v e g r o w t h r a t e s w e r e c l o s e l y c o r r e l a t e d w i t hl e a f a r e a p a r t i t i o n i n g i n s e v e n o f t h e n i n e s p e c i e s , b u t w e r e i n v e r s e l yc o r r e l a t e d w i t h l e a f w e i g h t p a r t i t i o n i n g f o r s i x o f t h e n i n e s p e c i e s .R e l a t i v e g r o w t h r a t e s w e r e p o o r l y c o r r e l a t e d w i t h n e t a s s i m i l a t i o n r a t e sf o r f i v e o f t h e n i n e s p e c i e s . T h e p r o d u c t o f n e t a s s i m i l a t i o n r a t e t i m e sl e a f a r e a p a r t i t i o n i n g i s s h o w n t o b e e q u a l t o t h e r e l a t i v e l e a f a r e ae x p a n s i o n r a t e .

T h e s e r e s u l t s i n d i c a t e t h a t g r o w t h r e s p o n s e s d u e t o t e m p e r a t u r e s h i f t sw e r e m o r e s e n s i t i v e t o c h a n g e s i n l e a f a r e a p a r t i t i o n i n g o r r e l a t i v e l e a fa r e a e x p a n s i o n r a t e s t h a n t o n e t a s s i m i l a t i o n r a t e s . B e c a u s e c h a n g e s i nl e a f a r e a p a r t i t i o n i n g o r r e l a t i v e l e a f a r e a e x p a n s i o n r a t e s c a n h a v e a ne f f e c t o n r e l a t i v e g r o w t h r a t e s t h a t o v e r s h a d o w c h a n g e s i n n e t a s s m i l a -t i o n r a t e s , a n d b e c a u s e n e t a s s i m i l a t i o n r a t e s a r e l a r g e l y a f u n c t i o n o fu n i t a r e a r a t e s o f p h o t o s y n t h e s i s , t h e c o r r e l a t i o n o f u n i t a r e a r a t e s o fp h o t o s y n t h e s i s w i t h g r o w t h s h o u l d i n c l u d e c o n s i d e r a t i o n o f l e a f a r e ap a r t i t i o n i n g o r r e l a t i v e l e a f a r e a e x p a n s i o n r a t e s .

F o r more t h a n a d e c a d e , c o n s i d e r a b l e e f f o r t h a s been e x -p e n d e d i n t h e s e a r ch f o r crop v a r i e t i e s or e c o t y p e s t h a t have h i g hu n i t l e a f ar e a p h o t o s y n t h e t i c r a t e s 3 - 6 , 1 1 , 1 3 , 1 4 , 1 6 , 2 2 ) .T h e s e i n v e s t i g a t i o n s h a v e been b a s e d on t h e a s s u m p t i o n t h a th i g h u n i t area p h o t o s y n t h e t i c r a t e s s h o u l d c o r r e l a t e w e l l w i t hg r o w t h . G e n e r a l l y , i t h a s been found t h a t g r o w t h i s not w e l lc o rr el a t e d w i t h u n i t a r ea r a t e s o f p h o t o s y n t h e s i s 4 , 1 3 , 1 5 , 1 6 , 8 2 3 ) , a l t h o u g h some a u t h o r s have r e p o r t e d otherwise 9 ) .H o w e v e r , g o o d c o r r e l a t i o n s between g r o w t h a n d r a t e s of l e a fa r ea e x p a n s i o n 5 - 7 , 1 1 , 2 2 , 2 4 , 2 6 ) have b e e n r e p o r t e d .As e a r l y a s 1 9 3 8 , Heath a n d G r e g o r y 1 2 ) concluded t h a t

P r e s e n t a d d r e s s : Ornamental P l a n t s Research L a b o r a t o r y , A g r i c u l -t u r a l R e s e a r c h S e r v i c e , U n i t e d S t a t e s D e p a r t m e n t of A g r i c u l t u re , Cor-v a l l i s , O r e g o n 9 7 3 3 0 .2 M i s s i s s i p p i A g r i c u l t u r a l a n d F o r e s t r y E x p e r i m e n t S t a t i o n c o o p e r a t -i n g .

r a t e s o f l e a f a r e a e x p a n s i o n h a v e a g r e a t e r i n f l u e n c e o n d r ym a t t e r p r o d u c t i o n t h a n d o n e t a s s i m i l a t i o n r a t e s . The NAR3 i sp r i m a r i l y a f u n c t i o n o f p h o t o s y n t h e s i s , p a r t i c u l a r l y d u r i n g t h ef i r s t f e w w e e k s f o l l o w i n g s e e d l i n g e m e r g e n c e . E x t e n s i v e s t u d i e sl a t e r v e r i f i e d t h e p r e d o m i n a n t r o l e o f l e a f a r e a e x p a n s i o n o v e rNAR a s a f a c t o r i n p l a n t g r o w t h 2 5 , 2 6 ) . H o w e v e r , a s E v a n s 8 )p o i n t s o u t , two r e c e n t d e v e l o p m e n t s h a v e r e n e w e d t h e i n t e r e s ti n t h e r o l e o f p h o t o s y n t h e s i s i n p l a n t g r o w t h . F i r s t , IR g a sa n a l y s i s h a s p e r m i t t e d t h e a c c u r a t e a n d r a p i d m e a s u r e m e n t o fs h o r t t e r m p h o t o s y n t h e t i c r a t e s . S e c o n d , t h e d i s c o v e r y t h a t t h ep a t h w a y s a n d r a t e s o f p h o t o s y n t h e s i s d i f f e r g r e a t l y among s p e -c i e s h a s l e d p l a n t s c i e n t i s t s t o r e - e x a m i n e t h e r o l e o f p h o t o s y n -t h e s i s i n p l a n t g r o w t h . C o u p l e d w i t h t h i s r e n e w e d i n t e r e s t i np h o t o s y n t h e s i s i s a g r o w i n g a w a r e n e s s t h a t p a r t i t i o n i n g o f p h o t o -s y n t h a t e i n t o n ew l e a f g r o w t h i s a l s o a n i m p o r t a n t f a c t o r i n p l a n tg r o w t h 2 , 1 0 , 1 7 , 1 9 , 2 1 ) . T h i s p a p e r i s a n a s s e s s m e n t o f t h er e l a t i o n b e t w e e n g r o w t h , NAR, a n d t h e p a r t i t i o n i n g o f d a i l yw e i g h t g a i n i n t o new l e a f a r e a a n d n ew l e a f w e i g h t .MATERIALS AND METHODS

N i n e s p e c i e s m a i z e , Z e a mays L . v a r . T r u c k e r s F a v o r i t e ;c o t t o n , G o s s y p i u m h i r s u t u m L . v a r . S t o n e v i l l e 2 1 3 ; s o y b e a n ,G l y c i n e m a x [ L . ] M e r r . v a r . P i c k e t t 7 1 ; v e l v e t l e a f , A b u t i l o nt h e o p h r a s t i M e d i c . ; s p u r r e d a n o d a , Anoda c r i s t a t a [ L . ] S c h l e c h t ;p r i c k l y s i d a , S i d a s p i n o s a L . ; common c o c k l e b u r , X a n t h i u mp e n s y l v a n i c u m W a l l r . ; j o h n s o n g r a s s , S o r g h u m h a l e p e n s e [ L . ]P e r s . ; r e d r o o t p i g w e e d , A m a r a n t h u s r e t r o f l e x u s L . ) w e r e g r o w nf r o m s e e d i n 1 5 - c m - d i a m e t e r p o t s o f s o i l i n g r o w t h c h a m b e r su n d e r t h r e e d a y / n i g h t t e m p e r a t u r e s 2 1 / 1 0 , 3 2 / 2 1 , a n d 3 8 /2 7 C ) . Chamber c o n d i t i o n s w e r e 7 0 + 5 r e l a t i v e h u m i d i t y ,w i t h 1 0 0 w m - 2 o f p h o t o s y n t h e t i c a l l y a c t i v e r a d i a t i o n a t t h ep l a n t c a n o p y t o p f o r 1 4 h r e a c h d a y . L e a f a r e a s w e r e m e a s u r e dw i t h a H a y a s h i Denko a u t o m a t i c a r e a m e t e r m o d e l AAM-54 Y e n E n t e r p r i s e s , I n c . , 2 9 1 2 T e r m i n a l T o w e r , C l e v e l a n d , O h i o4 4 1 1 3 ) , a n d d r y w e i g h t s o f l e a v e s , s t e m s , a n d r o o t s o f two p l a n t so f e a c h s p e c i e s w e r e d e t e r m i n e d t h r e e t i m e s a week f o r 4 w e e k sa f t e r e m e r g e n c e o r u n t i l t h e t o t a l p l a n t w e i g h t e x c e e d e d 5 g . Thep l a n t m a t e r i a l w a s d r i e d a t 7 0 C u n t i l i t r e a c h e d c o n s t a n t w e i g h t .D a t a w e r e a v e r a g e d f o r t h e t w o i n d i v i d u a l s o f e a c h s p e c i e sh a r v e s t e d a t o n e t i m e .3 A b b r e v i a t i o n s : NAR: n e t a s s i m i l a t i o n r a t e ; L A P : l e a f a r e a p a r t i t i o nc o e f f i c i e n t ; LWP: l e a f w e i g h t p a r t i t i o n c o e f f i c i e n t ; LAR: l e a f a r e a r a t i o ;k , : r e l a t i v e g r o w t h r a t e ; k I : r e l a t i v e l e a f a r e a e x p a n s i o n r a t e ; k L : r e l a t i v el e a f w ei g h t g r o w t h r a t e .4 M e n t i o n o f a t r a d e m a r k o r p r o p r i e t a r y p ro d u c t d o e s n o t c o n s t i t u t e ag u a r a n t e e o r w a r r a n t y o f t h e p r o d u c t b y t h e U . S . D e p a r t m e n t o fA g r i c u l t u r e o r t h e M i s s i s s i p p i A g r i c u l t u r a l a n d F o r e s t r y E x p e r i m e n tS t a t i o n , a n d d o e s n o t i m p l y i t s a p p r o v a l t o t h e e x c l u s i o n o f o t h e rp r o d u c t s t h a t ma y a l s o b e s u i t a b l e .

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LEAF AREA PARTITIONING AND GROWTHT h e f o l l o w i n g e q u a t i o n s wer e f i t t o d a t a f o r l ea f a r e a e x p a n -s i o n , l e a f w e i g h t i n c r e a s e , and t o t a l w e i g h t i n c r e a s e s u m o f l e a f ,s t e m , a n d r o o t w e i g h t ) 2 , 2 8 ) :

A = A o e k a (1 )w h e r e A : l e a f area a t a n y g i v e n t i m e d M 2 ) ; A O : e x t r a p o l a t e dv a l u e o f l e a f area on d a y o f e m e r g e n c e a t t i m e z e r o ) ; k 5 : r e l a t i v el e a f ar e a e x p a n s i o n r a t e / d a y ; t : t i m e d a y s ) .

W = W o e w t 2 )w h e r e W: t o t a l w e i g h t a t a n y g i v e n t i m e m g ) ; W 0 : e x t r a p o l a t e dv a l u e o f t o t a l w e i g h t on d a y o f e m e r g e n c e a t t i m e z e r o ) ; k w :r e l a t i v e g r o w t h r a t e / d a y .

L = L o e k L t 3 )w h e r e L : l e a f w e i g h t a t a n y g i ve n t i me m g ) ; L:e x t r a p o l a t e dv a l u e o f l e a f w e i g h t on d a y o f e merg e n ce a t t i m e z e r o ) ; k L :r e l a t i v e l e a f w e i g h t g r o w t h r a t e / d a y . The l e a f a rea p a r t i t i o nc o e f f i c i e n t i s d e f i n e d as

LP dA/dtd W / d tI f e q u a t i o n s 1 , 2 , a n d 3 d e s c r i b e g r o w t h ,

k A 0 e k sAP w.e~tb e c a u s e i f

y=bekx d y = k b e k Xdx

S i m i l a r l y , t h e l e a f w e i g h t p a r t i t i o n c o e f f i c i e n t i s d e f i n e d a sLWPL / d tdW/dt

a n d c a l c u l a t e d a skLLoekLtLWP k w W o e k w t x

N e t a s s i m i l a t i o n r a t e s (NAR = d W x 2 8 ) were c a l c u l a t e da s

- k w W o e kAoeksa

T h e v a l u e f o r t u s e d i n t h e c a l c u l a t i o n o f LAP, LWP, a n d NARwa s t h e t i m e a t t h e m i d p o i n t o f t h e r e g r e s s i o n l i n e s d e s c r i b e d b ye q u a t i o n s 1 , 2 , a n d 3 . The m i d p o i n t t i m e wa s o n e - h a l f o f t h e 2 8-d a y g ro w t h p e r i o d o r o n e - h a l f o f t h e t i m e r e q u i r e d f o r t h eo b s e r v e d t o t a l d r y w e i g h t t o r e a c h 5 g , w h i c h e v e r o c c u r r e d f i r s t .

RESULTSE q u a t i o n s 1 , 2 , a n d 3 f i t t h e d a t a f o r a r e a , t o t a l w e i g h t , a n dl e a f w e i g h t , r e s p e c t i v e l y . Of t h e 8 1 c o r r e l a t i o n c o e f f i c i e n t s o fl o g , A , l o g , W, or l o g , L versus t i m e , 5 9 were 0 . 9 8 o r g r e a t e r ,a n d o n l y s i x were l e s s t h a n 0 . 9 5 . None wa s l e s s t h a n 0 . 9 1 .T h e e f f e c t o f t e m p e r a t u r e on r e l a t i v e g r o w t h r a t e s k w ) , r e l a -t i v e l e a f area e x p a n s i o n r a t e s k 5 ) , a n d r e l a t i v e l e a f w e i g h tg r o w t h r a t e s k L ) i s s h o w n i n T a b l e I I n a l l c a s e s , t h e v a l u e s o fk w , k a n d k L were g r e a t e s t a t 3 2 / 2 1 a n d l o w e s t a t 2 1 / 1 0 C . T h ev a l u e s o f k w a n d k . were s t r o n g l y c o r r e l a t e d f o r a l l s p e c i e s a t a l lt e m p e r a t u r e s F i g . 1 ) . T h e p o i n t s l a y s c a t t e r e d c l o s e t o a l i n ed e s c r i b e d b y k a = k w , w h i c h i n d i c a t e s t h a t , f o r t h e s e s p e c i e s , t h e

r a t e s o f r e l a t i v e l e a f area e x p a n s i o n a n d r e l a t i v e g r o w t h a r en e a r l y e q u a l .To p e r m i t t h e r e a d e r t o c a l c u l a t e t h e v a l u e s o f LAP, NAR,a n d LWP, t h e v a l u e s f o r A O , W0 a n d L o f o r e a c h s p e c i e s a n dt e m p e r a t u r e r e g i m e a re g i v e n i n T a b l e I I As i n t h e ca s e o f t h e kv a l u e s , t h e s e i n t e r c e p t s c h a n g e d i n r e s p o n s e t o t e m p e r a t u r es h i f t s . I n c o n t r a s t t o t h e k v a l u e s , w h e r e i n c r e a s e s i n k w werea s s o c i a t e d w i t h i n c r e a s e s i n b o t h k . a n d k L , i n c r e a s e s i n k w weren o t w e l l c o r r e l a t e d w i t h c h a n g e s i n W 0 , A O, or L o .D u r i n g e x p o n e n t i a l g r o w t h , k 5 , k w , a n d k L a re c o n s t a n t s .H o w e v e r , LAP, NAR, a n d LWP c a n c h a n g e w i t h t i m e s o onemust s p e c i f y t h e a g e o f t h e p l a n t when r e f e r r i n g t o any p a r t i c u l a rv a l u e o f LAP, NAR, or LWP. The v a l u e s f o r LAP, NAR, a n dLWP f o r a l l s p e c i e s a t e a c h temperature were c a l c u l a t e d u s i n gt h e t i m e a t t h e m i d p o i n t o f t h e g r o w t h p e r i o d T a b l e I I ) a n dwere p l o t t e d versus k w F i g s . 2 , 3 , a n d 4 ) . F i g u r e 2 s h o w s t h a tf o r a l l s p e c i e s e x c e p t m a i z e a n d c o c k l e b u r , e a c h i n c r e a s e i n LAPwas a s s o c i a t e d w i t h an i n c r e a s e o f a b o u t t h e same r e l a t i v e s i z e i nk w . F o r c o c k l e b u r , t h e LAP o f t h e p l a n t s grown a t 3 8 / 2 7 wass l i g h t l y g r e a t e r t h a n t h a t o f t h e p l a n t s grown a t 3 2 / 2 1 , e v e nt h o u g h t h e k w o f t he 3 2 / 2 1 p l a n t s g r e a t l y e x c e e d e d t h a t o f t h e3 8 / 2 7 p l a n t s .I n c o n t r a s t , t h e c o r r e l a t i o n b e t w e e n NAR a nd k w was notn e a r l y as g o o d a s t h e c o r r e l a t i o n b e t w e e n LAP a n d k w F i g . 3 ) .F o r p r i c k l y s i d a , v e l v e t l e a f , j o h n s o n g r a s s , p i g w e e d , a n d s p u r r e da n o d a , l a r g e c h a n g e s i n k w were a s s o c i a t e d w i t h l i t t l e or noc h a n g e i n NAR. M o r e o v e r , f o r s o y b e a n , v e l v e t l e a f , s p u r r e d

Table I. Relative Growth Rates , R e l at iv e Le a f A re a Expansion Rates k 5 ) , a n d Relati v e LeefWeight Growth Rates k L) f o r Nine Species Grown U n d er Three Temperature RegimesTh e k val u e s have u n its of time-1, an d are slopes of expone ntial growth equations that i n c l u d ed a ta f rom 0 to 28 d a y s . If th e total d r y w e ig ht of a single i n d i v i d u a l of a species e x c e e de d 5 gprior to 28 days, harvesting was terminated an d the slopes an d intercepts were d etermi n ed f ro m d a taobtaine d u p to that time.

2 1 / 1 0 C 3 2 / 2 1 C 38/27 CS p e c i e s k w k a k1 k . k a kL k v k a kLM a i z e 0 .096 0.133 0.159 0.255 0.354 0.376 0.178 0.189 0.206Cotton 0 . 0 8 6 0.073 0.0 83 0 .206 0.197 0.20 0 0 .1 88 0.172 0 .1 88S o y b e a n 0 . 1 0 8 0.124 0.107 0.202 0.199 0.195 0.165 0.168 0.159V e l v e t l e a f 0 . 1 4 2 0.153 0.158 0.294 0. 289 0.297 0.249 0.260 0.270S p u r r e d a n o d a 0 . 1 3 5 0.131 0 .1 39 0.253 0.260 0.268 0.234 0 .244 0.255Prickly s id a 0.11 4 0.098 0.085 0.268 0.274 0.286 0.246 0.223 0.241C o c k l e b u r 0 . 1 6 5 0.151 0.161 0.269 0.263 0.252 0.204 0.176 0.1 89J o h n s o n g r a s s 0 . 1 5 6 0 .1 39 0.156 0.391 0.370 0.421 0 .359 0.324 0.358P i g w e e d 0 . 2 6 2 0.239 0.267 0.482 0.436 0.463 0.393 0.328 0.382

1 1l a n t P h y s i o l V o l . 5 9 , 1 9 7 7

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1 2 POTTER AND JONESa n o d a , p r i c k l y s i d a , j o h n s o n g r a s s , a n d p i g w e e d , t h e m aximu mNAR v a l u es d i d n o t c o r r e s p o n d w i t h t h e m aximu m k v a l u e s .P a r t i t i o n i n g o f t o t a l d a i l y w e i g h t g a i n i n t o new l e a f w e i g h t(LWP) w a s p o o r l y c o r r e l a t e d w i t h k W F i g . 4 ) . F o r m a i z e , c o t -t o n , s o y b e a n , s p u r r e d a n o d a , a n d c o c k l e b u r , a g e n e r a l d e c r e a s ei n LWP w a s a s s o c i a t e d w i t h a n i n c r e a s e i n k w . I n c o n t r a s t , ap o s i t i v e r e l a t i o n b e t w e e n LWP a n d k w w a s o b s e r v e d f o r t h eo t h e r s p e c i e s .

DISCUSSIONS e v e r a l i n v e s t i g a t o r s h a v e d e s c r i b e d t h e c l o s e c o r r e l a t i o n b e -t w e e n t h e r a t e o f l e a f a r e a e x p a n s i o n a n d g r o w t h 5 - 7 , 1 1 , 1 2 ,

2 2, 2 5, 2 6 ) . H o w e v e r , t h i s c o r r e l a t i o n d o e s n o t e s t a b l i s h a c a u s ea n d e f f e c t r e l a t i o n b e t w e e n l e a f a r e a e x p a n s i o n a n d g r o w t h r a t e .By a n a l y z i n g t h e i n v e s t m e n t o f d a i l y w e i g h t g a i n i n t o d a i l y l e a fa r e a e x p a n s i o n , o n e c a n e s t a b l i s h a c a u s e a n d e f f e c t r e l a t i o nb e t w e e n l e a f a r e a e x p a n s i o n a n d g r o w t h . T h e d a i l y g r o w t h o f ap l a n t i s t h e p r o d u c t o f NAR a n d a r e a . T h e p a r t i t i o n i n g o f d a i l yg r o w t h i n t o l e a f a r e a f o r t h e n e x t d a y w i l l d e t e r m i n e t h e a r e a o nt h e n e x t d a y a n d , h e n c e , t h e d a i l y g r o w t h o n t h e n e x t d a y w i l ld e p e n d t o a l a r g e e x t e n t o n t h e v a l u e o f t h e LAP.T h e LAP i s d i f f e r e n t f r o m t h e LAR 2 8 ) . LAR i s t h e l e a f a r e ad i v i d e d b y t h e t o t a l p l a n t w e i g h t a t a n y g i v e n t i me , w h e r e a s LAPi s t h e d a i l y c h a n g e i n LAR. T h e t w o p a r a m e t e r s d i f f e r f u n d a -m e n t a l l y b e c a u s e NAR x LAR = k w 2 8 ) ; b u t NAR x LAP =

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0 0 . 1 0 . 2 0 . 3 0 . 4 0 .5R E L A T I V E LEAF AREA E X P A N S I O N RATE d m - d 6 2 1 d a 0 1 )F I G . 1 . R e l a t i v e g r o w t h r a t e s k w ) v e r s u s r e l a t i v e l e a f a r e a e x p a n s i o nr a t e s k a ) f o r n i n e s p e c i e s g r o w n a t t h r e e t e m p e r a t u r e r e g i m e s . F o r e a c hs p e c i e s , t h e v a l u e s o f b o t h k w a n d k . w e r e h i g h e s t a t 3 2 / 2 1 a n d l o w e s t a t2 1 / 1 0 C .

P l a n t P h y s i o l . V o l . 5 9 , 1 9 7 7k a T h e v a l u e o f LAP i s t h a t i t s h o w s t h e p h y s i o l o g i c a l r e l a t i o n -s h i p b e t w e e n NAR a n d k a . A l s o , LAP h a s m o r e p h y s i o l o g i c a lm e a n i n g t h a n e i t h e r k a o r LAR, b e c a u s e LAP c o n t a i n s i n f o r m a -t i o n a b o u t t h e r a t e a n d d i s p o s i t i o n o f d r y m a t t e r p ro d u c t i o n a sw e l l a s t h e r a t e o f l e a f a r e a e x p a n s i o n .D u r i n g e x p o n e n t i a l g r o w t h , k w , k a , a n d k L a r e c o n s t a n t s , b u tLAP a n d NAR w i l l c h a n g e w i t h t i m e u n l e s s k w = k a . S i m i l a r l y ,LWP w i l l c h a n g e w i t h t i m e u n l e s s k w = k L . A s t e p w i s e s o l u t i o no f t h e LAP e q u a t i o n g i v e s

LAP = kaAmkwwmw h e r e Am a n d W m a r e v a l u e s f o r a r e a a n d w e i g h t c a l c u l a t e d a st h e m i d p o i n t o f t h e g r o w t h p e r i o d . Am a n d Wm a r e p o i n t s o n t h er e g r e s s i o n l i n e s a n d a r e n o t d a t a p o i n t s . S i n c e t h e c o n f i d e n c el i m i t s a b o u t a r e g r e s s i o n l i n e a r e n a r r o w e s t a t t h e m i d p o i n t o ft h e l i n e , t h e c o n f i d e n c e i n LAP c a l c u l a t e d a t t h e m i d p o i n t i sg r e a t e s t . T h i s r a t i o n a l e a p p l i e s t o t h e c a l c u l a t i o n o f NAR a n dLWP u s i n g m i d p o i n t t i m e s .T h e p o o r c o r r e l a t i o n b e t w e e n NAR a n d g r o w t h h a s b e e n w e l ld o c u m e n t e d 1 2 , 2 7 ) . More r e c e n t l y , n u m e r o u s i n v e s t i g a t o r sh a v e r e p o r t e d t h a t u n i t a r e a p h o t o s y n t h e t i c r a t e s d o n o t c o r r e -l a t e w e l l w i t h g r o w t h 5 - 7 , 1 1 , 2 2 , 2 3 ) . S i n c e t h e u n i t a r e a r a t eo f p h o t o s y n t h e s i s i s t h e p r i m a r y c o m p o n e n t o f NAR, i t i s n o ts u r p r i s i n g t h a t t h e r e l a t i o n s h i p s b e t w e e n g r o w t h a n d NAR a n db e t w e e n g r o w t h a n d u n i t a r e a r a t e s o f p h o t o s y n t h e s i s a r e s i m i -l a r .

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T a b l e II . Y I n t e r c e p t Values f o r T o t a l D r y W e i g h t W O ) . L e a f Area A O ) , a n d L e a f W e i g h t L O ) f o r N i n eS p e c i e s Grown Under Three T e m p e r a t u r e R e g i m e sTime at t h e m i d p o i n t wa s o n e - h a l f t h e duration o f t h e g r o w t h p e r i o d .

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W O A O L 0 m i d p o i n tS p e c i e s m g ) d m 2 ) m g ) d a y s ) m g ) d m 2 ) mg ) d a y s ) m g ) d m 2 ) m g ) d a y s )Maize 1 6 5 0 . 0 8 3 1 7 . 8 1 3 7 1 . 5 0 . 0 3 7 7 . 4 6 6 1 2 8 0 . 1 2 2 9 . 5 1 0Cotton 6 3 . 5 0 . 0 7 7 4 3 . 1 1 4 5 2 . 6 0 . 0 8 1 3 2 . 2 1 2 5 0 . 5 0 . 0 9 0 2 8 . 6 1 4S o y b e a n 9 4 . 8 0.070 5 1 . 6 1 4 6 8 . 3 9 0 . 1 0 3 7 . 6 1 2 7 7 . 5 0.081 4 2 . 0 1 3V e l v e t l e a f 7 . 4 9 0 . 0 0 5 7 2 . 3 0 1 4 5 . 9 3 0 . 0 1 1 2 . 8 9 1 3 8 . 8 2 0 . 0 1 0 2 . 7 8 1 4S p u r r e d a n o d a 8 . 5 4 0 . 0 0 7 3 4 . 2 5 1 4 7 . 0 2 0 . 0 0 8 9 2 . 6 4 1 4 6 . 1 2 0.0068 1 . 9 4 1 4P r i c k l y sida 2 . 6 8 0.0031 2 . 2 1 1 4 1 . 4 8 0 . 0 0 2 6 0 . 5 7 1 4 2 . 3 0 . 0 0 5 1 1 . 1 8 1 4Cocklebur 3 0 . 5 0 . 0 4 7 1 8 . 0 1 4 2 1 . 2 0 . 0 3 1 1 3 . 7 1 1 2 7 . 1 0 . 0 6 2 1 8 . 9 1 4J o h n s o n g r a s s 5 . 5 7 0 . 0 0 5 6 1 . 4 5 1 4 3 . 6 6 0.0091 0 . 9 1 1 0 3 . 0 8 0 .0 0 9 1 1 . 3 2 1 1P i g w e e d 0 . 2 6 0 . 0 0 0 6 6 0.11 1 4 0 . 2 6 0 . 0 0 1 3 0 . 1 9 4 1 2 0 .4 3 0 . 0 0 2 3 0 . 3 1 1 4

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LEAF AREA PARTITIONING AND GROWTH

NET ASSIMILATION RATE m g r d m d a y )F I G . 3 . R e l a t i v e g r o w t h r a t e s k w ) c o m p a r e d w i t h n e t a s s i m i l a t i o n

r a t e s o f n i n e s p e c i e s grown a t t h r e e t e m p e r a t u r e r e g i m e s . F o r e a c hs p e c i e s , t h e k w v a l u e s were h i g h e s t a t 3 2 / 2 1 a n d l o w e s t a t 2 1 / 1 0 C .-b

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(g - d a y ~ g d a )F I G . 4 . R e l a t i v e g r o w t h r a t e s k w ) c o m p a r e d w i t h l e a f w e i g h t p a r t i -t i o n i n g o f n i n e s p e c i e s grown a t t h r e e t e m p e r a t u r e r e g i m e s . F o r e a c hs p e c i e s , t h e k w v a l u e s were h i g h e s t a t 3 2 / 2 1 a n d l o w e s t a t 2 1 / 1 0 C .D i f f e r e n c e s i n g r o w t h among s p e c i e s i n t h i s s t u d y were d u e t od i f f e r e n t i a l responses t o t e m p e r a t u r e . The o b s e r v a t i o n t h a t f o r

s e v e r a l s p e c i e s j o h n s o n g r a s s , p i g w e e d , s p u r r e d a n o d a , a n dp r i c k l y s i d a ) , n e t a s s i m i l a t i o n r a t e s were n e a r l y i n d e p e n d e n t o ft e m p e r a t u r e may h a v e b e e n d u e t o t h e r e l a t i v e l y l o w l e v e l o fp h o t o s y n t h e t i c a l l y a c t i v e r a d i a t i o n 1 0 0 w-m-2) i n t h e growthc h a m b er s . T h i s l a c k o f a temperature e f f e c t on NAR was n ot d uet o t h e p a t h w a y o f CO 2 f i x a t i o n , b e c a u s e j o h n s o n g r a s s a nd p i g -weed a re d i c a r b o x y l i n a c i d s p e c i e s , whereas s p u r r e d ano da a n dp r i c k l y s i d a a re C a l v i n c y c l e s p e c i e s . I n a l l f o u r s p e c i e s , NARv a l u e s c h a n g e d l i t t l e d u e t o temperature s h i f t s .

I n t h i s s t u d y , LAP was m u c h b e t t e r c o r r e l a t e d with growtht h a n was NAR. T h e s e d a t a i n d i c a t e t h a t l e a f area p a r t i t i o nc o e f f i c i e n t s a re e x t r e m e l y s e n s i t i v e to temperature, a n d theys u g g e s t t h a t g r o w t h r e s p o n s e s t o temperature i n t h i s s t u d y werel a r g e l y d u e t o d i f f e r e n c e s i n LAP. The t e m p e r a t u r e e f f e c t onLAP was c o n s i s t e n t w i t h M i l t h o r p e s 2 0 ) c o n c l u s i o n t h a t l e a farea e x p a n s i o n i s s t r o n g l y i n f l u e n c e d by t e m p e r a t u r e . Also,t h e s e d a t a d e m o n s t r a t e t h a t i n t h e a b s e n c e o f m a r k e d d i f f e r e n c e si n NAR, g r o w t h can b e d r a s t i c a l l y a f f e c t e d by cha n ges i n LAP.F i n a l l y , t h e s e d a t a suggest a reason f o r t h e f a i l u r e of NAR andp h o t o s y n t h e s i s m e a s u r e m e n t s to c o r r e l a t e w e l l with growth. I tseems r e a s o n a b l e t h a t t h e r e l a t i v e growth rate o f a s p e c i e s couldbe e n h a n c e d i f t r a i t s f o r h i g h rates o f p h o t o s y n t h e s i s were com-b i n e d w i t h t r a i t s f o r h i g h l e a f area p a r t i t i o n c o e f f i c i e n t s .I f NAR i s c o n s t a n t , a c h a n g e i n LAP l e a d s to a proportion a lc h a n g e i n k a . Taken i n c o n j u n c t i o n with u n i t area rates ofp h o t o s y n t h e s i s , i n f o r m a t i o n about e i t h e r LAP or ka i s e q u a l l yu s e f u l f r o m t h e s t a n d p o i n t o f combining t r a i t s to increase rela-

t i v e g r o w t h r a t e s . LAP i s u s e f u l i n t h i s c o n t e x t i n t h a t i t i s ameasure o f t h e process t h a t a l l o w s two p l a n t s w i t h d i f f e r e n tNAR v a l u e s t o h a v e t h e s a m e v a l u e o f k a

B e c a u s e LAP, NAR, a n d LW P a r e c a l c u l a t e d f r o m i n f o r m a -t i o n a b o u t A O , W 0 , L o k k w , a n d k L , t h e r e are two p o i n t s a b o u tt h e s e s l o p e a n d i n t e r c e p t v a l u e s t h a t n e e d e m p h a s i s . F i r s t , A O ,W0, a n d L o were d e t e r m i n e d b y e x t r a p o l a t i n g r e g r e s s i o n l i n e s t ot i m e zero e m e r g e n c e ) . The i n s t a n t o f e m erg e n c e i s d i f f i c u l t t od e t e r m i n e a n d t h i s l e a d s t o errors i n t h e i n t e r c e p t v a l u e s . A l s o ,s e e d l i n g s r e q u i r e some t i m e a f t e r e m erg e n c e b e f o r e t h e y r e a c hf u l l p h o t o s y n t h e t i c competence. One w o u l d n o t e x p e c t t h eg r o w t h o f s e e d l i n g s d u r i n g t h e f i r s t 2 or 3 d a y s a f t e r e merg e n cet o c o i n c i d e w i t h t h e e x p o n e n t i a l g r o w t h f u n c t i o n s t h a t f i t t h eg r o w t h d a t a f r o m 3 t o 2 8 d a y s a f t e r emergence. T h u s , A O , W 0 ,a nd L o may not b e t h e a c t u a l a r e a or w e i g h t s a t emergence, a n dA O , W0 , a nd L o may c h a n g e w i t h f a c t o r s t h a t a l t e r processeso c c u r r i n g p r i o r t o emergence or t h e s l o p e s o f t h e e x p o n e n t i a lg r o w t h f u n c t i o n s . S e c o n d l y , t h e c l o s e c o r r e l a t i o n b e t w e e n k a a n dk w i s m i s l e a d i n g . W h i l e k a / k w was a p p r o x i m a t e l y 1 . 0 f o r mosts p e c i e s a t t h e t h r e e temperatures, t h e d e p a r t u r e s f r o m u n i t y a rea m p l i f i e d b e c a u s e LAP a n d NAR a re e x p o n e n t i a l f u n c t i o n s o f k aa nd k w .Th e e x p r e s s i o n s u s e d t o c a l c u l a t e LAP a n d NAR i n d i c a t e t h a tLAP i s i n v e r s e l y r e l a t e d t o k w a n d NAR i s d i r e c t l y r e l a t e d t o k wwhen k a =k w . The o b s e r v a t i o n t h a t i n res pon s e t o t e m p e r a t u r es h i f t s , LAP i s b e t t e r c o r r e l a t e d w i t h k w t h a n wa s NAR i s d u e t ot h e s i m u l t a n e o u s c h a n g e s i n k a , k w , A O , a n d W0 a s s o c i a t e d w i t htemperature s h i f t s . S i m i l a r l y , t h e s i mu l t a n e o u s c h a n g e s i n k L ,k w , L o , a n d W 0 combined t o m ak e LWP p o o r l y c o o r r e l a t e d w i t hk w .J a c k s o n 1 9 ) c o i n e d a term c a l l e d t h e l e a f - t o - t o t a l - g r o w t hr a t i o , w h i c h i s i d e n t i c a l t o LWP. H e s u g g e s t e d t h a t a f a c t o r t h a ti n c r e a s e d t h e l e a f - t o - t o t a l - g r o w t h r a t i o c o u l d i n c r e a s e t h e r e l a -t i v e g r o w t h rate w i t h o u t i n c r e a s i n g t h e net a s s i m i l a t i o n r a t e .However, t h e mere i n v e s t m e n t o f d r y matter p r o d u c t i o n i n t ol e a f w e i g h t i s a poor i n d e x o f p o t e n t i a l g r o w t h , a s e v i d e n c e d b yt h e f a i l u r e o f l e a f w e i g h t p a r t i t i o n i n g t o c o r r e l a t e w e l l w i t hg r o w t h . I n f a c t , f o r s e v e r a l s p e c i e s m a i z e , c o t t o n , s o y b e a n ,s p u r r e d a n o d a , an d c o c k l e b u r ) , t h e v a l u e s o f l e a f w e i g h t p a r t i -t i o n i n g were h i g h e s t a t t h e l o w temperature, w h e r e b o t h k w a n dk a were m i n i m a l .

B l a c k e t a l . 1 ) s u g g e s t e d t h a t p l a n t s t h a t h a v e h i g h u n i t l e a farea r a t e s o f p h o t o s y n t h e s i s a re l i k e l y t o b e strong c o m p e t i t o r s i nc r o p p i n g s i t u a t i o n s . T h i s work e x t e n d s t h a t o f B l a c k e t a l b yi n d i c a t i n g t h a t t h e c o m p e t i t i v e a b i l i t y o f any s p e c i e s , r e g a r d l e s so f u n i t l e a f area p h o t o s y n t h e t i c r a t e , i s enhanced by an i n c r e a s ei n t h e l e a f area p a r t i t i o n c o e f f i c i e n t .

Acknowkdgment-We are g r a t e f u l to J . D. Hesketh fo r s t i m u l a t i n g our t h i n k i n g a nd f o rp o i n t i n g ou t many sources of p e r t i n e n t l i t e r a t u r e . We thank R. D . Wauchope f o r suggesting th eformat f o r t h e graphs a n d f o r many h e l p f u l d i s c u s s i o n s , a n d weth ank E. L. John son f o r h i se x c e l l e n t t e c h n i c a l a s s i s t a n c e .

LITERATURE CITED1 . BLACK, C . C . , T. M CHEN, AND R. H . BROWN. 1 9 69 . B io ch em i ca l basis fo r p l a n t

c o m p e t i t i o n . Weed S c i . 1 7 : 3 3 8 - 3 4 4 .2 . BLACKMAN, V . H . 1 91 9. The compound i n t e r e s t la w i n p l a n t growth. Ann. Bot. 33 : 353-

3 6 0 .3 . C A I L S O N , G. E . , R. B. P E A R C E , D. R. L E E , ANDR. H. HArr. 1 9 7 1 . P h o t o s y nt h e s i s and

p h o t o r e s p i r a t i o n i n two c l o n e s of orcha rdg ra s s. C r o p S c i . 11 : 3 5 - 3 7 .4 . C u t n s , P . E., W. L. OGREN, AN D R. H. HAGEMAN. 1969. V a r i e t a l effects in soybean

p h o t o s y n t h e s i s a n d photorespiration. C r o p S c i . 9 : 323-327.5. DELANEY, R. H.ANDA. K. DOBDENZ. 1974. Yield of a lf a lf a asrelated tocarbon exchange.

Agron. J. 6 6 : 49 8-5 00.6 . DUNCAN, W. G. ANDJ . D. H E s K E T H . 1 9 6 8 . Ne t photosynthetic rates, r e la t i ve leaf growth

r a t e s , a n d l e a f n u m b e r s of 2 2 races of maize grown at e i g h t temperatures. Crop. S ci . 8:670-674.

7 . EL-SHA V A w y , M., J. HESKETH, ANDH. M U R A M O r o . 1965. Leaf photosyntheticratesando t h e r growth ch aracteristics among26 species of Gossypium. Crop Sci. 5: 173-175.

8 . EVANS, L. T. 1975. Crops and world food supply,cropevolution and the origins ofphysiology. In : L. T. E v a n s , ed., Crop Physiology. Cambridge U n i v e r s i t y Press, GreatB r i t a i n . pp. 1 - 2 2 .

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1 4 POTTER AND JONES9 . F O U S O V A , S . AND N . A V R A T O V S C U K O V A . 1 9 6 7 . H y b r i d v i g o u r a n d p h o t o s y n t h e t i c r a t e o fl e a f d i s k s i n Z e a m a y s L . P h o t o s y n t h e t i c a 1 : 3 - 1 2 .1 0 . H AC K ET T , C . AND H . M. R A W S O N . 1 9 7 4 . An e x p l o r a t i o n o f t he c a r b o n e c o n o m y o f t h e

t o b a c c o p l a n t . I I . P a t t e r n s o f l e a f g r o w t h a n d d ry m a t t e r p a r t i t i o n i n g . A u s t . J . P l a n tP h y s i o l . 1 : 2 7 1 - 2 8 1 .

1 1 . H A N S O N , W. D . 1 9 7 1 . S e l e c t i o n f o r d i f f e r e n t i a l p r o d u c t i v i t y among j u v e n i l e m a i z e p l a n t s :a s s o c i a t e d n et p h o t o s y n t h e t i c r a t e a n d l e a f a r e a c h a n g e s . C r o p S c i . 1 1 : 3 3 4 - 3 3 9 .1 2 . H E A T H , 0 . V . S . AND F . G . G R E G OR Y . 1 9 3 8 . T h e c o n s t a n c y o f t h e mean n et a s s i m i l a t i o nr a t e a n d i t s e c o l o g i c a l i m p o r t a n c e . A n n . B o t . N . S . 2 : 8 1 1 - 8 1 8 .

1 3 . H E I C H E L , G . H . AND R . B . M U S G R A V E . 1 9 6 9 . V a r i e t a l d i f f e r e n c e s i n n et p h o t o s y n t h e s i s o fZ e a m a y s L . C r o p S c i . 9 : 4 8 3 - 4 8 6 .

1 4 . I R V I N E , J . E . 1 9 6 7 . P h o t o s y n t h e s i s i n s u g a r c a n e v a r i e t i e s u n d e r f i e l d c o n d i t i o n s . C r o p S c i .7 : 2 9 7 - 3 0 0 .

1 5 . I R V I N E , J . E . 1 9 7 5 . T h e r e l a t i o n s o f p h o t o s y n t h e t i c r a t e s a n d l e a f a n d c a n o p y c h a r a c t e r s t os u r g a r c a n e y i e l d . C r o p S c i . 1 5 : 6 7 1 - 6 7 6 .1 6 . I Z H A R , S . AND D . H . W A L L A C E . 1 9 6 7 . S t u d i e s o n t h e p h y s i o l o g i c a l b a s i s o f y i e l d d i f f e r e n c e s .I I I . G e n e t i c v a r i a t i o n i n p h o t o s y n t h e t i c e f f i c i e n c y o f P h a s e o l u s v u l g a r i s L. C r o p S c i . 7 :4 5 7 - 4 6 0 .

1 7 . K U M U R A , A . 1 9 7 5 . D r y m a t t e r p a r t i t i o n a n d c l i m a t i c f a c t o r s . I n : Y . M u r a t a , e d . , C r o pp r o d u c t i v i t y a n d s o l a r e n e r g y u t i l i z a t i o n i n v a r i o u s c l i m a t e s i n J a p a n . J I B P S y n t h e s i s V o l1 1 . U n i v e r s i t y o f Tokyo P r e s s .1 8 . McNAUGHTON, S . J . 1 9 7 4 . D e v e l o p m e n t a l c o n t r o l o f n e t p r o d u c t i v i t y i n T y p h a l a t i f o l i ae c o t y p e s . E c o l o g y 5 5 : 8 6 4 - 8 6 9 .

P l a n t P h y s i o l . V o l . 5 9 , 1971 9 . J A C K S o N , J. E . 1 96 3 . R e l a t i o n s h i p o f r e l a t i v e l e a f g r o w t h r a t e t o n et a s s i m i l a t i o n r a t e a n d i

r e l e v a n c e t o t h e p h y s i o l o g i c a l a n a l y s i s o f p l a n t y i e l d . N a t u r e 2 0 0 : 9 0 9 .2 0 . M I L T H O R P E , F . L. 1 9 5 9 . S t u d i e s o n t h e e x p a n s i o n o f t h e l e a f s u r f a c e . I The i n fl u en c e o

t e m p e r a t u r e . J . E x p . B o t . 1 0 : 2 3 3 - 2 4 9 .2 1 . M O N S I , M. AND Y . M U R A T A . 1 9 6 9 . D e v e l o p m e n t o f p h o t o s y n t h e t i c s y s t e m s a s i n f l u e n c e

b y d i s t r i b u t i o n o f m a t t e r . I n : P r e d i c t i o n a n d M e a s u r e m e n t o f P h o t o sy n t h et i c P r o d u c t i v i t yP r o c e e d i n g s o f t h e I B P / P P T e c h n i c a l M e e t i n g , T r e b o n , C z e ch o s l o v ak i a . pp . 1 1 5 - 1 2 9 .

2 2 . MURAMOTO, H. , J . H E S K E T H , AND M. E L - S H A R K A W Y . 1 9 6 5 . R e la t i on s h ip s among r a t el e a f a r e a d e v e l o p m e n t , p h o t o s y n t h e t i c r a t e , a n d r a t e o f d ry m a t t er p ro d u ct i o n amonA m e r i c a n c u l t i v a t e d c o t t o n s a n d o t h e r s p e c i e s . C r o p S c i . 5 : 1 6 3 - 1 6 6 .

2 3 . S H I B L E S , R . M. AND H . A . MAcDONALD. 1 9 6 2 . P h o t o s y n t h e t i c a r e a a n d r a t e i n r e l a t i o n ts e e d l i n g v i g o r o f b i r d s f o o t t r e f o i l L o t u s c o r n i c u l a t u s L . ) . C r o p S c i . 2 : 2 9 9 - 3 0 2 .2 4 . S L A T Y E R , R . 0 . 1 9 7 0 . C o m p a r a t i v e p h o t o s y n t h e s i s , g r o w t h a n d t r a n s p i r a t i o n o f t w o s p e c io f A t r i p l e x . P l a n t a 9 3 : 1 7 5 - 1 8 9 .2 5 . W A T S O N , D . J . 1 9 4 7 . C o m p a r a t i v e p h y s i o l o g i c a l s t u d i e s o n t h e g r o w t h o f f i e l d c r o p s .

V a r i a t i o n i n n e t a s s i m i l a t i o n r a t e a n d l e a f a r e a b e t w e e n s p e c i e s a n d v a r i e t i e s , a n d w i t h ia n d b e t w e en y e a r s . Ann. B o t . N . S . 1 1 : 4 1 - 7 6 .2 6 . W A T S O N , D . J . 1 9 4 7 . C o m p a r a t i v e p h y s i o l o g i c a l s t u d i e s o n t h e g r o w t h o f f i e l d c r o p s . I I . T he f f e c t o f v a r y i n g n u t r i e n t s u p p l y o n n e t a s s i m i l a t i o n r a t e a n d l e a f a r e a . Ann. B o t . N . S . 13 7 5 - 4 0 7 .

2 7 . W A T S O N , D . J . 1 9 5 2 . P h y s i o l o g i c a l b a s i s o f v a r i a t i o n i n y i e l d . A d v . A g r o n . 4 : 1 0 1 - 1 4 5 .2 8 . W E S T , C . , G . E . B R I G G S , AND F . K I D D . 1 9 2 0 . M e t h o d s a n d s i g n i f i c a n t r e l a t i o n s i n t hq u a n t a t i v e a n a l y s i s o f p l a n t g r o w t h . Ne w P h y t o l . 1 9 : 2 0 0 - 2 0 7 .


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