Sugarcane Physiology

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Sugarcane physiology

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  • P H Y S I O L O G Y O F S U G A R - C A N E

    V I I . E F F E C T S O F T E M P E R A T U R E , P H O T O P E R I O D D U R A T I O N , A N D D I U R N A L A N D

    S E A S O N A L T E M P E R A T U R E C H A N G E S O N G R O W T H A N D R I P E N I N G

    B y K . T . G L A S Z I O U , * T . A . B U L L , * M . D . H A T C H , * a n d P . C . W H I T E M A N * t

    [ M a n u s c r i p t r e c e i v e d J u l y 2 7 , 1 9 6 4 ]

    S u m m a r y

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

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

    u n d e r c o n t r o l l e d e n v i r o n m e n t s . D u r i n g t h e f i r s t 3 m o n t h s o f g r o w t h , d a y a n d n i g h t

    t e m p e r a t u r e e f f e c t s w e r e m a i n l y a d d i t i v e , b u t a t 6 m o n t h s t h e i n t e r a c t i o n e f f e c t s

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

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

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

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

    a t t h e o p t i m u m t e m p e r a t u r e ( 3 0 C ) f o r d r y m a t t e r p r o d u c t i o n . S u g a r c o n c e n t r a t i o n

    d i d n o t e x c e e d 1 2 % f r e s h w e i g h t w h e n t e m p e r a t u r e s w e r e m a i n t a i n e d c o n s t a n t

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

    o n a r e l a t i v e l y l o n g - t e r m b a s i s , h i g h s u g a r l e v e l s ( 1 7 % o f f r e s h w e i g h t ) w e r e a t t a i n e d

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

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

    s u g a r - c a n e .

    1 . I N T R O D U C T I O N

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

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

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

    ( v a n D i l l e w i j n 1 9 5 2 ) . H o w e v e r , c o n f l i c t i n g r e s u l t s o b t a i n e d b y d i f f e r e n t w o r k e r s

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

    F i e l d d a t a f r o m T a i w a n ( S u n a n d C h o w 1 9 4 9 ) a n d H a w a i i ( C l e m e n t s , S h i g e u r a ,

    a n d A k a m i n e 1 9 5 2 ) s h o w a p o s i t i v 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 s t e m e l o n g a t i o n

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

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

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

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

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

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

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

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

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

    r e s p o n s e o f t h e p l a n t t o e x t e r n a l f a c t o r s d e c r e a s e s w i t h a g e ( B o r d e n 1 9 4 5 ) . S h a w

    ( 1 9 5 3 ) a n a l y s e d d a t a f r o m a l a r g e n u m b e r o f c a n e - g r o w i n g a r e a s a n d f o u n d l i t t l e

    * D a v i d N o r t h P l a n t R e s e a r c h C e n t r e , T h e C o l o n i a l S u g a r R e f i n i n g C o . L t d . , T o o w o n g ,

    B r i s b a n e .

    t P r e s e n t a d d r e s s : B o t a n y D e p a r t m e n t , H e b r e w U n i v e r s i t y , I s r a e l .

    A u s t . J . B i o l . S c i . , 1 9 6 5 , 1 8 , 5 3 - 6 6

  • 54 K. T. GLASZIOU ET AL.

    correlation between the sucrose concentration in cane juice and the ratio of summer to winter temperature when the ratio was between 10 and 1'3, while above 13 the

    TABLE 1 ANALYSIS OF VARIANCE FOR DAY AND NIGHT TEMPERATURE EFFECTS ON DRY WEIGHT PRODUCTION, STALK LENGTH, NODE NUMBER, TILLER NUMBER AND WEIGHT, AND STALK DIAMETER AT 106-DAY

    HARVEST INTERVAL Experiment 1

    F Ratios

    Source of Degrees

    Variation of Total Stalk Stalk Leaf Node Tiller Tiller Stalk Freedom Dry Length Dry Dry No. No. Fresh Dia-Weight Weight Weight Weight meter ---------------------

    Day temperature (T) 4 22'9** 38'9** 48'1** 42** 171** 16'3** 6'1** 7'2**

    Night tempera-ture (t) 4 10'7** 12'6** 9'5** 8'6** 25'8** n.s. n.s. 8'8**

    Txt interaction 16 n.st 2'83** n.S. n.S. n.s. n.s. 1'8* n.S.

    Error~ 75 0033 0035 0116 0018 128 133 336 0056 ----------

    * P < 005. ** P < 001. t n.s., not significant. ~ Values in this line are error mean squares.

    TABLE 2 ANALYSI8 OF VARIANCE FOR DAY AND NIGHT TEMPERATURE EFFECTS ON DRY WEIGHT PRODUCTION, STALK LENGTH, NODE NUMBER, TILLER NUMBER AND WEIGHT, AND STALK DIAMETER AT 20!)-DAY

    HARVEST INTERVAL Experiment 1

    F'Ratios

    Source of Degrees

    Variation of Total Stalk Stalk Leaf Node Tiller Tiller Stalk Freedom Dry Length Dry Dry No. No. Dry Dia-Weight Weight Weight Weight meter ------------------

    Day temperature (T) 4 9'0** 16'4** 12'7** 12'6** 14'9** n.st 3'8* 3'6**

    Night tempera-ture (t) 4 4-7* 6'1** 5'7** 82** 32'2** 2'8* n.s. n.s.

    Txt interaction 16 9-1** 2'0* 6'0** 50** 4'5** n.s. 4'0** 4'2**

    Error~ 125 0011 0005 004 0006 298 144 328 0-058 * P < 005. ** P < 001. t n.s., not significant.

    ~ Values in this line are error mean squares.

    sugar content fell. These results are contrary to the notion that low temperature promotes sucrose content. Furthermore there was no consistent relationship between

  • P H Y S I O L O G Y O F S U G A R - C A N E . V I I 5 5

    T A B L E 3

    A N A L Y S I S O F V A R I A N C E F O R D A Y A N D N I G H T T E M P E R A T U R E E F F E C T S O N S U G A R C O N T E N T O F

    S T A L K S A N D L E A V E S

    E x p e r i m e n t 1

    F R a t i o s

    S o u r c e o f V a r i a t i o n

    D e g r e e s o f

    S u g a r i n S t a l k s

    S u g a r i n L e a v e s

    F r e e d o m

    l O 6 D a y s

    2 0 9 D a y s 1 0 6 D a y s

    2 0 9 D a y s

    D a y t e m p e r a t u r e ( T ) 4

    2 0 ' 5 * *

    2 6 ' 3 * * 8 ' 7 * * 3 3 ' 3 * *

    N i g h t t e m p e r a t u r e ( t ) 4

    4 ' 7 *

    n . s t

    n . s . 4 2 *

    T x t i n t e r a c t i o n 1 6

    3 ' 1 * *

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    n . s .

    E r r o r : j :

    2 5 1 5 9 9

    0 ' 0 0 l 5

    3 8

    0 0 0 0 2

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    : j : V a l u e s i n t h i s l i n e a r e e r r o r m e a n s q u a r e s .

    T A B L E 4

    A N A L Y S I S O F V A R I A N C E F O R D A Y A N D N I G H T T E M P E R A T U R E , P H O T O P E R I O D D U R A T I O N , A N D D I U R N A L

    T E M P E R A T U R E C Y C L E E F F E C T S O N D R Y W E I G H T P R O D U C T I O N , S T A L K L E N G T H , N O D E N U M B E R ,

    S T A L K D I A M E T E R , T O T A L S U G A R P E R P L A N T , A N D S U G A R P E R 1 0 0 G F R E S H W E I G H T O F P R I M A R Y

    S o u r c e o f

    V a r i a t i o n

    D a y t e m p e r a t u r e ( T )

    N i g h t t e m p e r a t u r e ( t )

    P h o t o p e r i o d

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    c y c l e ( h )

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    S T A L K S A T 2 0 0 - D A Y H A R V E S T I N T E R V A L

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    n . s .

    1 0 5

    1 9 4 6 3 4 1 1 0 0 4

    * P < 0 0 5 . * * P < 0 0 l . t n . s . , n o t s i g n i f i c a n t .

    S u g a r

    p e r

    P l a n t

    n . s .

    n . s .

    1 6 ' 4 * *

    n . S .

    n . s .

    3 ' 3 *

    3 2 ' 4 * *

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    n . s .

    2 3 3

    : j : O n l y i n t e r a c t i o n s f o r w h i c h s i g n i f i c a n t e f f e c t s w e r e o b t a i n e d a r e s h o w n .

    V a l u e s i n t h i s l i n e a r e m e a n s q u a r e s .

    I I 3 6 d e g r e e s o f f r e e d o m .

    S t a l k

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  • 56 K. T. GLASZIOU ET AL.

    sucrose content and the altitude of the growing area ranging from sea level to 1300 ft for 53 areas in Mexico. Shaw concluded that sugar concentration is an inherent func-tion of latitude, reaching peak values at 18 north and south latitudes, and suggested that the causal agency was the day length during the critical period of crop maturation rather than temperature per se.

    MAIN EFFECTS OF DAY AND NIGHT TEMPERATURE INTERACTION OF DAY AND NIGHT TEMPERATURE - 209

    '8 106 DAYS 2-4 209 DAYS .!2. ~~'>., (p~~~,)l :: .!2. l-

    I- X

    i:5 ',7

    "'[ ~ > > is ~ ',6 !ie ,20 I- !z ~ '5 l/ z "' "' ~ "5 it 2" (p~~~,)l -' LSD I g 34 g (P

  • P H Y S I O L O G Y O F S U G A R - C A N E . V I I

    5 7

    I I . M A T E R I A L S A N D M E T H O D S

    ( a ) G r o w t h C o n d i t i o n s

    U n i f o r m o n e - b u d s e t s f r o m t o p i n t e r n o d e s o f c v . P i n d a r w e r e g e r m i n a t e d a n d

    g r o w n i n v e r m i c u l i t e i n 2 - g a l l o n c o n t a i n e r s a n d w a t e r e d d a i l y , o n c e w i t h c o m p l e t e

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

    f o u r t o e a c h t r u c k . G r o w t h o f a l l t r e a t m e n t s , w h i c h c o m m e n c e d i n M a r c h 1 9 6 2 ,

    M A I N E F F E C T S O F D A Y A N D N I G H T T E M P E R A T U R E

    2 3

    2 1

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    T E M P E R A T U R E ( O c )

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    I N T E R A C T I O N O F D A Y A N D N I G H T T E M P E R A T U R E - 2 0 9 D A Y S

    (p~~'~l)I

    3 4

    F i g . 3 . - E f f e c t s o f d a y a n d n i g h t t e m p e r a t u r e o n s t a l k d r y w e i g h t p r o d u c t i o n . E x p e r i m e n t 1 .

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

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

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

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

    o f t r u c k s .

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    T h e p h y t o t r o n s y s t e m u s e d i s d e s c r i b e d i n t h e 1 9 6 4 A n n u a l R e p o r t o f t h i s

    I n s t i t u t i o n a n d i s e s s e n t i a l l y t h e s a m e a s i n t h e E a r h e a r t P h y t o t r o n ( W e n t 1 9 5 7 ) .

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

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

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

  • 58 K. T. GLASZIOU ET AL.

    material. Confirmation of this assumption was provided by the similarity of error mean square terms obtained at the 106- and 209-day harvests of experiment 1 as the latter harvest involved six replicate plants, three on each of two trucks. Lack of significance of the third-order interaction term in experiment 2 also supported the basic assumption as it provided an estimate of between-truck variation.

    1-0

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    MAIN EFFECTS OF DAY AND NIGHT TEMPERATURE

    106 DAYS

    (p~~-~,)I

    I ! I 18 22 26 30 34

    TEMPERATURE (Oc)

    1-4

    ~ 1"3 '1l z ',2 w

    " :i? .101 g

    1-0

    209 DAYS

    DAY TEMPERATURE

    NIGHT TEMPERATURE

    (p~~~,)I I! I! 18 22 26 30 34

    TEMPERATURE (Oc)

    INTERACTION OF DAY AND NIGHT TEMPERATURE - 209 DAYS

    ; 25[ ~ 15 ~ w 5 Q 34 o z

    !) 36 ~r

    )'("-11,0 26"

    ~.-?4rU,.lj) 22

  • P H Y S I O L O G Y O F S U G A R - C A N E . V I I 5 9

    ( d ) S a m p l i n g M e t h o d s

    F r e s h w e i g h t o f t h e w h o l e p l a n t ( i n c l u d i n g t i l l e r s b u t e x c l u d i n g r o o t s ) w a s

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

    M A I N E F F E C T S O F D A Y A N D N I G H T

    T E M P E R A T U R E

    M A I N E F F E C T S O F D A Y T E M P E R A T U R E I N T E R A C T I O N O F D A Y A N D N I G H T T E M P E R A T U R E - 2 0 9 D A Y S

    1 0 6 D A Y S

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  • 60 K. T. GLASZIOU ET AL.

    Mean internode length was calculated by dividing stalk length by node number minus 1. Mean stem diameter was also measured.

    The plant parts were oven-dried at 70C, reweighed, and immediately ground in a 5-in. laboratory mill. The ground samples were sealed in plastic bags and stored for sugar analysis.

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    d a y a n d n i g h t t e m p e r a t u r e s f r o m 1 8 t o 3 0 0 . W h e r e a p p l i c a b l e t h e s e t r a n s f o r m a t i o n s

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

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    p r e s e n t e d i n g r a p h i c a l f o r m o n l y w h e n t h e s e a t t a i n e d s i g n i f i c a n c e a t t h e 1 0 % l e v e l .

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

    o n d a t a a r e p r e s e n t e d i n T a b l e s 1 - 3 f o r e x p e r i m e n t 1 , a n d i n T a b l e 4 f o r e x p e r i m e n t 2 .

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    t h e c o a s t a l r i v e r s o f N e w G u i n e a ( B r a n d e s 1 9 5 8 ) . I n t h e s e a r e a s t h e a v e r a g e m a x i m u m

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    a b o v e 3 4 0 o r f a l l b e l o w 2 2 0 . S e a s o n a l v a r i a t i o n s a r e s m a l l b e t w e e n t h e c o o l e r a n d

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    3 0 m i n b e t w e e n t h e e x t r e m e t i m e s o f s u n r i s e a n d s u n s e t d u r i n g t h e c o u r s e o f t h e y e a r .

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    a c t i o n e f f e c t s o f 1 2 - h o u r l y c h a n g e s o f d a y a n d n i g h t t e m p e r a t u r e o n g r o w t h a n d

    d e v e l o p m e n t d u r i n g t h e f i r s t 3 m o n t h s ( T a b l e 1 ; F i g s . 1 - 8 ) . H o w e v e r , s i g n i f i c a n t

    i n t e r a c t i o n e f f e c t s w e r e o b s e r v e d f o r t h e n e x t 3 - m o n t h p e r i o d , m u c h o f w h i c h w a s

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

  • 62 K. T. GLASZIOU ET AL.

    particularly at 22, 26, and 300. The constant-temperature effect was much reduced when the photoperiod was decreased to 8 hr [Figs. 9, 10, 11, 12(a)], and except for certain combinations for which no general pattern is evident [Figs. 12(b), 12(c)], there was almost no effect of varying the photoperiod and thermoperiod cycles in or out of phase with one another. In view of the constancy ofthe natural environment where sugar-cane evolved, it is perhaps not surprising that it grows well under controlled constant-temperature regimes. However, this is in sharp contrast with many other plants which show a requirement for thermoperiodicity (Went 1961).

    The stalk is the storage organ for sugar in the sugar-cane plant. For illustrative purposes the time course of development of the stalk may be divided into four characteristic stages: germination, juvenile, and early and late adult stages, the latter being terminated either by onset of flowering or senescence.

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

  • 64 K. T. GLASZIOU ET AL.

    comparison of 106- and 209-day data show an apparent shift downwards in the optimum temperature for growth as the age of the plant increases (Figs. 1-3). This effect appears to be entirely due to the use of a chronological rather than a physio-logical time scale. For example stalk growth at 180 constant temperature compared with 300 gave a ratio of 0034 during the first 106-day interval, 023 during the second 103 days, and 109 during the next 81 days. For the last period, plants at 180 had just entered the early adult stage whereas those at 300 were in the late adult stage.

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

    o b s e r v e d , a c c o m p a n i e d b y a r a p i d d e c l i n e i n s u c r o s e c o n t e n t t o 6 5 % f r e s h w e i g h t

    a f t e r 3 5 d a y s . W h e n t h e s e p l a n t s w e r e r e t u r n e d t o 1 7 C c o n s t a n t t e m p e r a t u r e , s t a l k

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

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

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

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

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

    f o r 4 m o n t h s a t a n a v e r a g e t e m p e r a t u r e o f 2 6 C , t h e n t r a n s f e r r e d t o a c o n s t a n t -

    t e m p e r a t u r e g r e e n h o u s e a t 1 7 C . A f t e r 1 6 0 d a y s o f t h i s t r e a t m e n t , s o m e p l a n t s

    w e r e t r e a t e d f o r 3 5 d a y s a t 3 0 C , t h e n r e t u r n e d t o 1 7 C .

    W e s u g g e s t t h a t t h e p e a k v a l u e s f o r t h e s u c r o s e c o n c e n t r a t i o n i n c a n e j u i c e a t 1 8

    n o r t h a n d s o u t h l a t i t u d e s o b s e r v e d b y S h a w ( 1 9 5 3 ) a r e d u e t o a n o p t i m u m c o m -

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

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

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

    I V . A C K N O W L E D G M E N T S

    T h e a u t h o r s a r e i n d e b t e d t o D r . W . O . M c C a r t h y , U n i v e r s i t y o f Q u e e n s l a n d ,

    a n d M r . K . P . H a y d o c k , D i v i s i o n o f T r o p i c a l P a s t u r e s , C S I R O , f o r a s s i s t a n c e w i t h

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

    V . R E F E R E N C E S

    B O R D E N , R . J . ( 1 9 3 7 ) . - C a n e g r o w t h s t u d i e s . T h e e f f e c t o f s u n l i g h t o n t h e u t i l i z a t i o n o f n i t r o g e n

    a n d p o t a s h b y H I 0 9 c a n e . H a w a i i a n P l a n t e r s ' R e c . 4 1 : 3 - 5 .

    B O R D E N , R . J . ( 1 9 4 5 ) . - T h e e f f e c t o f n i t r o g e n f e r t i l i z a t i o n u p o n t h e y i e l d a n d c o m p o s i t i o n o f s u g a r

    c a n e . H a w a i i a n P l a n t e r s ' R e c . 4 9 : 2 5 9 - 3 1 2 .

    B R A N D E S , E . W . ( 1 9 5 8 ) . - " S u g a r c a n e . " ( U . S . D . A . A g r i c . H a n d b o o k N o . 1 2 2 . )

  • 66 K. T. GLASZIOU ET AL.

    CLEMENTS, H. F., SHIGEURA, G., and AKAMINE, E. K. (1952).-Factors affecting the growth of sugar cane. Tech. Bull. Univ. Hawaii Agric. Exp. Sta. No. 18.

    DILLEWIJN, C. VAN (1952).-"Botanyof Sugarcane." (Chronica Botanica Co.: Waltham, Mass.) HATCH, M. D., and GLASZIOU, K. T. (1963).-Sugar accumulation in sugar cane. II. Relationship

    of invertase activity to sugar content and growth rate in storage tissue of plants grown in controlled environment. Plant Physiol. 38: 344--8.

    MARTIN, J. P., and ECKART, R. C. (1933).-The effect of various intensities of light on the growth of the H109 variety of sugarcane. Hawaiian Planters' Rec. 37: 53-66.

    SHAW, H. R. (1953).-An international glance at sucrose content of cane. Proc.lnt. Soc. Sugarcane Technol. 8: 283--9.

    SUN, V. G., and CHOW, N. P. (1949).-The effect of climatic factors on the yield of sugar cane in Taiwan. Multiple factors investigation. Rep. Taiwan Sug. Exp. Sta. No.2. pp. 44--98.

    WENT, F. W. (1957).-ln "The Experimental Control of Plant Growth". (Chronica Botanica Co.: Waltham, Mass.)

    WENT, F. W. (1961).-ln "Encyclopedia of Plant Physiology". Vol. 16. pp. 1-23. (Ed. W. Ruhland.) (Springer-Verlag: Berlin.)

    WHITEMAN, P. C., BULL, T. A., and GLASZIOU, K. T. (1963).-The physiology of sugar-cane. VI. Effects of temperature, light, and water on set germination and early growth of Saccharum spp. Aust. J. Biol. Sci. 16: 416-28.

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