PHYSIOLOGY OF SUGAR-CANE

VII. EFFECTS OF TEMPERATURE, PHOTOPERIOD DURATION, AND DIURNAL AND

SEASONAL TEMPERATURE CHANGES ON GROWTH AND RIPENING
By K. T. GLASZIOU,* T. A. BULL,* M. D. HATCH,* and P. C. WHITEMAN*t
[Manuscript received July 27, 1964]
Summary
Independent and interaction effects of day and night temperature, photo-
period duration, and diurnal thermoperiodicity were studied on sugar-cane grown
under controlled environments. During the first 3 months of growth, day and night
temperature effects were mainly additive, but at 6 months the interaction effects
of all variables were numerous and complex. Many of the interaction effects could
be attributed to increased responses to constant-temperature regimes with a 12-hr
photoperiod. No evidence for thermoperiodicity requirements was found.
Sugar production per plant and sugar concentration in the stalk were highest
at the optimum temperature (30C) for dry matter production. Sugar concentration
did not exceed 12% fresh weight when temperatures were maintained constant
or when diurnal variations were given. However, when the temperature was varied
on a relatively long-term basis, high sugar levels (17% of fresh weight) were attained
by moving plants from high to low temperature. Sugar loss was incurred for the
reverse situation. The results are discussed in relation to the ripening process in
sugar-cane.
1. INTRODUCTION
Studies of temperature effects on growth and ripening of sugar-cane have given
rise to the concept that growth at high average temperatures is conducive to develop-
ment of a crop yielding a high weight of cane per acre but with low sugar content
(van Dillewijn 1952). However, conflicting results obtained by different workers
leave some doubt as to the validity of this generalization.
Field data from Taiwan (Sun and Chow 1949) and Hawaii (Clements, Shigeura,
and Akamine 1952) show a positive correlation between the rate of stem elongation
and both mean monthly minimum and maximum temperatures. For the Taiwan
data the first and second orders of partial correlation coefficients for response to mean
monthly minimum temperature were much larger than for mean monthly maximum
temperature, which could be taken to indicate that temperature during the dark
period is more important in controlling stem elongation. A negative correlation was
obtained between sugar content and mean monthly minimum temperature.
Interpretation of field results is complicated in that high average temperatures
are experienced when the day lengths are longest, and, for many areas, when the
rainfall is greatest: The stage of development of the crop also affects results as the
response of the plant to external factors decreases with age (Borden 1945). Shaw
(1953) analysed data from a large number of cane-growing areas and found little
* David North Plant Research Centre, The Colonial Sugar Refining Co. Ltd., Toowong,
Brisbane.
t Present address: Botany Department, Hebrew University, Israel.
Aust. J. Biol. Sci., 1965, 18, 53-66
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.
r r o r ~ 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**
r r o r ~ 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
PHYSIOLOGY OF SUGAR-CANE. VII 55
TABLE 3
ANALYSIS OF VARIANCE FOR DAY AND NIGHT TEMPERATURE EFFECTS ON SUGAR CONTENT OF
STALKS AND LEAVES
Experiment 1
F Ratios
Source of Variation
Degrees of
Sugar in Stalks Sugar in Leaves
Freedom
lO6 Days 209 Days 106 Days 209 Days
Day temperature (T) 4 20'5** 26'3** 8'7** 33'3**
Night temperature (t) 4 4'7* n.st
n.s. 42*
Txt interaction 16 3'1** n.s. 8'9** n.s.
Error:j: 25 1599 0'00l5 38 00002
* p < 005. ** P < 00l. t n.s., not significant.
:j: Values in this line are error mean squares.
TABLE 4
ANALYSIS OF VARIANCE FOR DAY AND NIGHT TEMPERATURE, PHOTOPERIOD DURATION, AND DIURNAL
TEMPERATURE CYCLE EFFECTS ON DRY WEIGHT PRODUCTION, STALK LENGTH, NODE NUMBER,
STALK DIAMETER, TOTAL SUGAR PER PLANT, AND SUGAR PER 100 G FRESH WEIGHT OF PRIMARY
Source of
Variation
Day temperature (T)
Night temperature (t)
Photoperiod
duration (H)
Diurnal temperature
cycle (h)
Interactions:j:
Txt
txHxh
TxHxt
TxHxh
TxHxtxh
Error
STALKS AT 200-DAY HARVEST INTERVAL
Experiment 2
F Ratios
Degrees
of Total
Stalk Node
Stalk
Freedom Dry
Length No.
Dia-
Weight meter
2 n.st n.s. 7'9** n.s.
1 n.s. 7'0* 32'9** n.s.
2 33'3** 10'5* 4'9* 28'9**
1 n.s. n.s. n.s. n.s.
4 n.s. n.s. n.s. n.s.
2 n.s. n.s. n.s. n.s.
4 13'0** 9'0* n.s. 7'7**
2 n.s. n.s. n.s. n.s.
4 n.s. 3'0* n.s. n.s.
105 1946 34 11 004
* P < 005. ** P < 00l. t n.s., not significant.
Sugar
per
Plant
n.s.
n.s.
16'4**
n.S.
n.s.
3'3*
32'4**
7'9**
n.s.
233
:j: Only interactions for which significant effects were obtained are shown.
Values in this line are mean squares.
II 36 degrees of freedom.
Stalk
Sugar
Concen-
tration
n.S.
n.s.
250**
n.s.
7'3*
n.s.
n.s.
n.s.
4'5**
027411
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.

::
.!2. l-
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X
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>
>
is
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LSD I g 34
g (P<D'D')
I- "4 oj
.,; .,;
9
9 9
'-3
,
..
zz z. 3D 3.
TEMPERATURE (Oc)
Fig. I.-Effects of day and night temperature on dry weight production. Experiment 1.
Shading experiments at the Hawaiian Sugar Planters' Experiment Station
(see Annual Report for 1961) have shown that growth of cane in the field increases
as the light intensity is increased up to full sunlight. Compared with shaded plants,
growth in full sunlight gave thicker but shorter stalks, broader and greener leaves,
and a greater rate of tiller production (Martin and Eckart 1933). Day length effects
have been demonstrated by Borden (1937) in that plants exposed all day to full
MAIN EFFECTS OF DAY AND NIGHT TEMPERATURE.

.!d-
'"
1'5
1'0
In
106 DAYS
DAY
TEMPERATURE

Q'SL ;2 2*6 30 3'4
TEMPERATURE ('c)
209 DAYS
a Z"r DAY iTEM/PERATURE NIGHT
"9 TEMPERATURE

"' tn "7

1'5 18 22 26 30 34
TEMPERATURE ('c)
INTERACTION OF DAY AND NIGHT TEMPERATURE -106 DAYS
"
i2[
-' ,
3
a
9 34
3D
Z.
"22
18 'IA\G'(..'\
Fig. 2.-Effects of day and night temperature on stalk length. Experiment 1.
sunlight produced more dry matter than plants exposed until noon or from noon to
sunset, the actual being affected by the amount of fertilizer applied. These
results show that light intensity and duration of the photoperiod must also be
considered when attempting to evaluate the effects of temperature on the plant.
The studies described in this paper were conducted under controlled environ-
mental conditions to establish with greater certainty the independent and interaction
effects of day and night temperatures and duration of photoperiod on growth,
development, and ripening of the cane plant.
PHYSIOLOGY OF SUGAR-CANE. VII 57
II. MATERIALS AND METHODS
(a) Growth Conditions
Uniform one-bud sets from top internodes of cv. Pindar were germinated and
grown in vermiculite in 2-gallon containers and watered daily, once with complete
nutrient solution and once with de-ionized water. The containers were placed on trucks,
four to each truck. Growth of all treatments, which commenced in March 1962,
MAIN EFFECTS OF DAY AND NIGHT TEMPERATURE
23
21
:
19

"7

"5
o

<5
g
1'1
106 DAYS
DAY
TEMPERATURE
L.S.D I
(P<OOl)
Q.gL '" ! .
18 22 26 30 34
TEMPERATURE (Oc)
2VG DAYS
22
:
j: 20
"

1-8
1-6
1
./j

1-4
'-2
22 26 30
TEMPERATURE (Oc)
INTERACTION OF DAY AND NIGHT TEMPERATURE - 209 DAYS

34
Fig. 3.-Effects of day and night temperature on stalk dry weight production. Experiment 1.
occurred under natural daylight in controlled environment greenhouses. All tem-
pe;rature and photoperiod changes were brought about by shifting trucks to the
appropriate greenhouse or constant-temperature darkroom. Differences due to
locations within a greenhouse were randomized by daily movements and rotation
of trucks.
1-7
1-6
-"2.

,,5

,,4


13
g
1-2
I-I
MAIN EFFECTS OF DAY AND NIGHT TEMPERATURE
106 DAYS
LSD. I
(P<O-Ol)
1 I I
18 22 26 30 34
TEMPERATURE (Oc)
,..

Cl ',7


<.3 ',6
g
1-5
209 DAYS
I I!
18 22 26 30 34
TEMPERATURE (Oc)
INTERACTION OF DAY AND NIGHT TEMPERATURE - 209 DAYS
:
2_O
t

>- "5
10 34
<5
g
L.S.D.
(P<O-Ol)'
Fig. 4.-Effects of day and night temperature on leaf dry weight production. Experiment 1.
The phytotron system used is described in the 1964 Annual Report of this
Institution and is essentially the same as in the Earheart Phytotron (Went 1957).
The use of four single plant replications grown on the same truck was considered
justified because of the high degree of environmental control, the randomization of
position effects, and the lack of genetic variation in asexually propagated planting
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
'"
og

0'8
:i?
g 0-7
0-6
MAIN EFFECTS OF DAY AND NIGHT TEMPERATURE
106 DAYS

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

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

)'("-11,0 26"
22
<" (0C) 18
Fig. 5.-Effects of day and night temperature on number of nodes per stalk. Experiment 1.
(b) Conditions for Experiment 1
Five day and five night temperatures from 18C to 34C at 4-degree intervals
were used in all combinations, temperature changes occurring at 12-hourly intervals.
There were four replicates per treatment for the first harvest interval at 106 days,
and six for the second at 209 days. A third harvest was made at 290 days on plants
grown at constant temperature only, with four replicates per treatment.
MAIN EFFECTS OF DAY TEMPERATURE
f- 4'0
z

30
:Q
j 2'0
i=
"-
o
1-0
'1l
z a
106 DAYS
40

U>
'"
30
='
f-
"-
20 0
f-
:0
(9
10


t::
INTERACTION OF DAY AND NIGHT TEMPERATURE - 209 DAYS
8T

:l 0 34
i=

-30
-26 ",,,,'1-"-

-22
'c) 18
Fig. 6.-Effects of day and night temperature on number and weight of tillers per
plant. Experiment 1.
(c) Conditions for Experiment 2
In this experiment plants were grown at all combinations of three day and-
night temperatures (22, 26, and 30C) with either an 8- or 12-hr photoperiod given
from 0800-1600 or 0600-1800 hr. Temperature changes were made on an 8/16 or
12/12 hourly cycle with changes at 0800 and 1600 hr or 0600 and 1800 hr, respectively.
Each treatment contained four replicates and plants were harvested after 200 days.
PHYSIOLOGY OF SUGAR-CANE. VII 59
(d) Sampling Methods
Fresh weight of the whole plant (including tillers but excluding roots) was
recorded, and then, individually, fresh weight of tillers, stem, and leaves plus leaf
MAIN EFFECTS OF DAY AND NIGHT
TEMPERATURE
MAIN EFFECTS OF DAY TEMPERATURE INTERACTION OF DAY AND NIGHT TEMPERATURE - 209 DAYS
106 DAYS 209 DAYS
11'
g
ffi 1'9
t;;
Z"f

o 1'7
"5!' !,
11'
g
Z'5
ffi 2'3

a 2"



1'9

DAY
TEMPERATURE
'2
c;
i 3[

1 34'
b 30

)'('41,0 26
18 22 26 3() 34 18 22 26 30
TEMPERATURE (Oc)
34
<tJi>4}; zt
TEMPERATURE (Oe)

(0C) 18
Fig. 7.-Effects of day and night temperature on diameter of stalks. Experiment 1.
-;:>
'"
"


>-
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"
30
20
z
o
>=
g

u
3 10
u
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'5
U1
40
t U1 20
0
:s 34
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if>
MAIN EFFECTS OF DAY AND NIGHT TEMPERATURE
106 DAYS 209 DAYS
I}"""
STALKS
I LS.D
q
LS.D
(P<O'OI)
I (P<O'OI)
" 40
-
z
30
r

20
o
u
LEAVES DAY
LEAVES
L.S.D
TEMPERATURE I (p<o.ol)
It! I r
'" 0: 10

TEMPERATURE

7' -0 ,LSD. TEMPERATURE
I (P<O'OI)
I I ! I
18 22 26 30 34 18 22 26 30 34
TEMPERATURE (OC) TEMPERATURE (OC)
INTERACTION OF DAY AND NIGHT TEMPERATURE - 106 DAYS
_ '3D
26 ,0',,'0
22

18
-;:>
'"
lOt
"
.'"
:; 5
0 l/
; 34
18
'26 30
,,'<-'0
'22

Fig. S.-Effects of day and night temperature on sugar concentration in stalks and leaves.
Experiment 1.
sheaths. Counting of nodes commenced with the pair encompassing an internode
exceeding 05 em in length. Stalk length was measured from base to apical meristem.
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.

I-
:r:
'"
300
200
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z
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o
o 14
z
MAIN EffECTS Of DAY AND NIGHT TEMPERATURE AND PHOTOPERIOD - 200 UAYS
P<O'Ol
PHOTOPERIOD (HR)
I
12
;/
12
PHOTOPERIOD (HR)
P<O'05
--------
'i'

:r:
I-
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zoo
ffi" 150
..J
:5
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t;;
100
-100

I-
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8 12.
PHOTOPERIOD (HR)
P<OOI

12
PHOTOPERIOD (HR)
P<OOI
/
'i'

:r:
I-
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200
z 150
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:5

100
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ffi
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Z
u
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.,
'"
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18
V>
'" o
o
z
::> L ::>
p<o'os
L
22 26 30
NIGHT TEMPERATURE ("c)
P<OOI
L_
8 12
PHOTOPERIOD (HR)
P<OOI
ci 14 I I , . 14L ,;:-'___ -= __ -,!
12 z 22 26 30 22 26 30
PHOTOPERIOD (HR) DAY TEMP;;:RATURE (Oc) NIGHT TEMPERATURE (Oc)
Fig. 9.-Main effects of day and night temperature and photoperiod duration on growth and
development of cane. Diurnal temperature cycles did not attain significance as a main effect
for any of the characters measured. Experiment 2.
(e) Sugar Analysis
The material from half of the replicates was bulked, prior to grinding, to give
two samples per treatment. Total sugars were determined in stem and leaf tissue.
The oven-dry material was extracted for 5 days with 70% ethanol, and the sugars
in the ethanolic extract determined as previously (Hatch and Glasziou 1963).
PHYSIOLOGY OF SUGAR-CANE. vn 61
(f) Data Processing
For experiment 1 at the first harvest interval at 106 days, transformation of
some of the measurements to logarithms shows approximately linear responses to
day and night temperatures from 18 to 300. Where applicable these transformations
have been made and for continuity have also been used for the later harvest interval.
All results were subjected to analysis of variance and the data presented are taken
from these analyses. In general, first- and second-order interaction effects have been
presented in graphical form only when these attained significance at the 10% level.
Significance of main effects and interactions was determined by testing against signific-
ant higher-order interactions containing the variables in question. Analyses of variance
on data are presented in Tables 1-3 for experiment 1, and in Table 4 for experiment 2.

....
:J:
" 4i
500
400
300
"
"
-'

....
o
....
200
100
INTERACTION OF DAY AND NIGHT TEMPERATURE AND PHOTOPERIOD -200 DAYS
P<OOl
12 - HR PHOTOPERIOD
...
.....
30

2.

22
",G

!,OO[
-' 100[
"
8
8 - HR PHOTOPERIOD
26
22

Fig. lO.-Interaction effects of day and night temperature and photoperiod duration on dry
weight production. Experiment 2.
III. RESULTS AND DISCUSSION
Sugar-cane is thought to have evolved on the flood plains and adjacent areas of
the coastal rivers of New Guinea (Brandes 1958). In these areas the average maximum
temperature is about 320 and the minimum about 230. Temperatures rarely rise
above 340 or fall below 220. Seasonal variations are small between the cooler and
warmer months, the range being about 3 degO, and there is only a difference of about
30 min between the extreme times of sunrise and sunset during the course of the year.
Our studies show that with a natural photoperiod, there are virtually no inter-
action effects of 12-hourly changes of day and night temperature on growth and
development during the first 3 months (Table 1; Figs. 1-8). However, significant
interaction effects were observed for the next 3-month period, much of which was
attributable to the favourable responses obtained at constant temperatures, and
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.
'i:
~
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w
"'
25
~ Z'O
o
'"


in
15
30
INTERACTION OF DAY AND NIGHT TEMPERATURE AND PHOTOPERIOD-ZOO DAYS
P<OOI
.... ,
'i:
~
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,...
w
25
~ 2'0
o
'"
~
15
8 - HR PHOTOPERIOD
D4
y
7lC 26
26 ('Q '26 ('(,1
""<:11
47UII
<: (OC)
"i?''-
22 , - ~ - ~
,!-\GY-
,,,,,,-
22 - ~ - ~
'!-\GY-'
Fig. H.-Interaction effects of day and night temperature and photoperiod duration on stalk
diameter. Experiment 2.
Environmental effects on the germination stage have been described previously
(Whiteman, Bull, and Glasziou 1963). Time courses for development of the primary
stalk at temperatures from 18 to 300 are shown in Figure 13. The shape of these
curves is mainly determined by the effect of temperature on the rate of increase of
leaf area and not on the rate of photosynthesis, since direct measurements made in
full sunlight show that photosynthesis per unit leaf area is virtually independent of
temperature over the range from 9 to 340 (Waldron and Glasziou, unpublished
data). During the juvenile stage the optimum temperature for plant and stalk
growth is about 300 (Figs. 1-3). As plants approach the early adult stage the leaf
area per stalk tends to a constant value, the rate of production of new leaves being
balanced by the senescence of old leaves. At this stage, the rate of stalk development
also tends to a constant value which is independent of temperature, but as the stalk
matures further the rate decreases. Thus stalks maintained at 220 enter early
adult stage at about 200 days, attaining a growth rate between 200 and 300 days
which is approximately equal to the maximum rate attained at 300 (Fig. 13). Direct
2
t-
Z
j
"-
150
100
'"
""
L?
::>
'"
50
g
/
30
/
/
/
PHYSIOLOGY OF SUGAR-CANE. VII
INTERACTION OF DAY AND NIGHT TEMPERATURE AND PHOTOPERIOD-ZOO DAYS
P<O'OI
...........
....
30
8
t-
Z


'"
""


6 - HR PHOTOPERIOD
D-1
y
26
"'''I;:I/

26 \2(,)

;':
g
26 \2(,)
22
",\",y..
I;: (OC)

22

INTERACTION OF DAY TEMPERATURE, PHOTOPERIOD, AND DIURNAL TEMPERATURE CYCLE -ZOO DAYS
P<O'OI
12 - HR PHOTOPERIOD 8 - HR PHOTOPERIOD
8
100
H
2
t-
Z
""
....

n
...
...
30
0

D-1).
;':
D-1 y 0
26 12
t-
26
<'11-1
6
<'11-1
(0 22

l'Ulrc- (0 22
C)

C)
INTERACTION OF NIGHT TEMPERATURE, PHOTOPERIOD, .AND DIURNAL TEMPERATURE CYCLE - 200 DAYS
P<005
8
100

'"
'" "-
50
g
'"

;':
g
12 - HR PHOTOPERIOD
30
8
!Z
n
0

8 - HR PHOTOPERIOD
g tvlGIf
26 12 t- 26 12
1;:"'''1;: '0 "'''1;:11 ,0'0
&0
III;: 22 6 0. I;: (0 ",0 ,
(DC) C)
63
Fig. 12.-Interaction effects of day and night temperatures, photoperiod duration, and diurnal
temperature cycle on total sugar per plant.
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.
For the two experiments so far reported the temperature and photoperiod
regimes constituting each treatment were maintained throughout the whole growth
period. Under such conditions, environments which gave the highest rate of dry
300
200
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TIME (DAYS)
Fig. 13.-Time course for growth of stalks at constant temperatures.
matter production also gave the most sugar per plant and the highest sugar content
on a dry weight basis (Figs. 1, 8,9, 12). The sucrose content reached a maximum value
in the stalk of about 12% fresh weight at 6 months of age and did not increase sub-
stantially over the next 3-month interval. Oontinuous growth at low temperature,
or with diurnal changes including high day and low night temperature, or the reverse,
did not result in higher sugar contents than obtained at 300 constant temperature.
An explanation for sugar contents of 12-20% of fresh weight often observed
in the field comes from studies in which plants raised in the field during the wet
summer season (average temperature 260) were transferred to controlled environ-
ments and watered well. A negative correlation was obtained between the rate of
stalk elongation and the rate of change of sugar content (Hatch and Glasziou 1963).
In a similar experiment the sucrose content of plants placed at 170 increased from
10 to 17% fresh weight during the first 90 days after which it declined slowly (Fig. 14).
PHYSIOLOGY OF SUGAR-CANE. VII 65
Some of the plants which had attained high sucrose level were transferred to 30C
constant temperature, upon which an exceedingly high rate of stalk elongation was
observed, accompanied by a rapid decline in sucrose content to 6 5% fresh weight
after 35 days. When these plants were returned to 17C constant temperature, stalk
elongation virtually ceased and sugar storage was resumed. Hence we reject the
notion that short-term (diurnal) variations in temperature are conducive to sugar
storage except that they be accompanied by an overall long-term change in average
temperature akin to the seasonal variations experienced under field conditions.
20
I- 16
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NON-SUGAR DRY WEIGHT
PLANTS KEPT AT 17C
PLANTS KEPT AT 30C
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4' I , , , ,
o 50 100 150 200 250
TIME (DAYS)
Fig. 14.-Changes in sugar contents of cane stalks in response to long-term
changes in temperature for growth. Plants were raised under field conditions
for 4 months at an average temperature of 26C, then transferred to a constant-
temperature greenhouse at 17C. After 160 days of this treatment, some plants
were treated for 35 days at 30C, then returned to 17C.
We suggest that the peak values for the sucrose concentration in cane juice at 18
north and south latitudes observed by Shaw (1953) are due to an optimum com-
bination of seasonal temperature and day-length variations. At latitudes closer to
the equator the temperature variation is too small, and at higher latitudes the day
lengths during the winter period are too short.
IV. ACKNOWLEDGMENTS
The authors are indebted to Dr. W.O. McCarthy, University of Queensland,
and Mr. K. P. Haydock, Division of Tropical Pastures, CSIRO, for assistance with
experiment design and statistical treatment of results.
V. REFERENCES
BORDEN, R. J. (1937).-Cane growth studies. The effect of sunlight on the utilization of nitrogen
and potash by HI09 cane. Hawaiian Planters' Rec. 41: 3-5.
BORDEN, R. J. (1945).-The effect of nitrogen fertilization upon the yield and composition of sugar
cane. Hawaiian Planters' Rec. 49: 259-312.
BRANDES, E. W. (1958).-"Sugarcane." (U.S.D.A. Agric. Handbook No. 122.)
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.