ELSEVIER

Field Crops Research 47 (1996) 143-153

Field
Crops
_
Research

Effect of nitrogen on the time-course of sucrose accumulation in

sugarcane
R.C. Muchow a,*, M.J. Robertson

b, A.W. Wood c, B.A. Keating a

a CSIRO Division of Tropical Crops and Pastures, St. Lucia, Queensland, Australia
b CS[RO Division of Tropical Crops and Pastures, Aitkenvale, Queensland, Australia
c CSR, Technical Field Department, lngham, Queensland, Australia

Received 4 December 1995; revised 28 February 1996; accepted 1 March 1996

Abstract
Sugarcane is harvested commercially at ages varying from 9 to 36 months. Since high N supply can decrease the sucrose
concentration in fresh millable stalks and consequently decrease the commercial value of the stalks, the opportunity exists to
manipulate N supply (both from fertiliser and that mineralised from soil organic matter) to maximise economic return at
different times of harvest. Accordingly, this study describes how N supply the time-course of sucrose accumulation in
sugarcane and determines yield both on a dry weight basis (as commonly analysed by crop physiologists) and on a fresh
weight basis (which is how cane is paid for commercially). Data on crop N uptake and its efficiency of utilisation are also
presented.
There was a trade-off between maximising sucrose yield and sucrose concentration in fresh millable stalks with different
N supply and this varied with time of harvest. Sucrose concentration in fresh millable stalks, particularly during early
growth, was maximised by low N supply. The lower sucrose concentration (on a fresh weight basis) with high N supply
could be largely explained by a decrease in stalk dry matter content. Most of the variation in stalk sucrose yield could be
explained by variation in stalk biomass irrespective of N supply. Whilst increasing N supply decreased the sucrose
concentration in dry millable stalks, this effect was relatively small compared to the large positive effect of N supply on stalk
biomass. It is concluded that N has a marked effect on stalk dry matter content, and hence a greater effect on the commercial
measures (yield and sucrose concentration of fresh millable stalks) of sugarcane production than on the physiological
measures (stalk biomass and sucrose concentration on a dry weight basis) of crop performance.
Keywords." Sugarcane; Nitrogen; Sucrose; Yield accumulation; N-use efficiency

1. Introduction
The m a n a g e m e n t of nitrogen (N) is central to
profitable and sustainable sugarcane production.
Yield limitations i m p o s e d by inadequate N supply

* Corresponding author.
0378-4290/96/$15.00 1996 Elsevier Science B.V.
Pll S 0 3 7 8 - 4 2 9 0 ( 9 6 ) 0 0 0 2 2 - 6

can have a large negative impact on profitability.

Conversely, N application in excess of crop requirem e n t m a y have potential deleterious off-farm impacts on the e n v i r o n m e n t and can also reduce sugar
quality and profitability. In commercial sugarcane
production, a p r e m i u m is paid to growers for high
sucrose concentration on a fresh weight basis. Num e r o u s studies have shown that high N application

144

R.C. Muchow et a l . / F i e l d Crops Research 47 (1996) 143-153

decreases sucrose concentration in fresh millable

stalks (Das, 1936; Borden, 1942, 1944, 1948; Hurney, 1984; Wood, 1990; Stevensen et al., 1992;
Chapman, 1994). Most of these studies have presented data at only one harvest time and have focused on fresh-weight measures of cane yield and
sucrose content. The few studies reporting timecourse responses were collected more than five
decades ago in Hawaii (Das, 1936; Borden, 1942,
1944, 1948). There is a need to confirm previous
responses with modern varieties and production systems. Sugarcane is harvested over a long period of
time. The timing of harvest is crucial to profitability
and this depends on the interaction between environment and the crop in relation to sucrose content and
on the economics of harvesting, transport and milling
operations. The opportunity exists to use N management to maximise profitability by manipulating the
sucrose concentration in fresh millable stalks at different crop ages. However, there is little information
published on the determinants of sucrose accumulation over time under variable N supply.
Payment to growers is based on stalk sucrose
yield calculated on the basis of fresh cane yield and
sucrose concentration on a fresh weight basis. The
use of fresh weight measures is important for commercial production, but this makes it difficult to
examine productivity responses in relation to driving
variables such .as N supply. For most agricultural
crops, biological yield is expressed on a dry weight
basis to overcome this problem in examining the
determinants of productivity. Similarly for sugarcane, variation in sucrose yield of millable stalks can
be examined in terms of stalk biomass and the
proportion of sucrose present in that biomass (sucrose
concentration g g ~ DW). Hence stalk dry matter
content is an important variable in relating biological
productivity to commercial yield in sugarcane production.
In this study, we report the growth of a plant crop
and a subsequent 1st ratoon crop of sugarcane grown
on the same site under high-input tropical conditions
and at three rates of N fertiliser application. The
objectives were (i) to examine how N affects the
time-course of yield accumulation in sugarcane; (ii)
to examine the partitioning of stalk biomass to sucrose in response to N supply; (iii) to examine the
change in stalk dry matter content over time in

response to N supply and the effect on the relationship between commercial and biological yield; and
(iv) to determine the efficiency of N utilisation in
biomass and sucrose production for the two crop
cycles.

2. Material and methods

2.1. Location and cultural details

The study was conducted on a fine sandy loam

(Macknade Series, (Wilson and Baker, 1990); USDA
Soil Taxonomy: Dystropept) at Macknade Research
Station, Ingham Queensland (18.7 S, 146.2 E). The
site had been under long-term sugarcane culture,
with the previous crop being harvested in 1991. The
experiment was managed to eliminate growth restrictions associated with water, soil and aerial pests and
diseases, and nutrient supply other than nitrogen.
The site was fumigated on 13 July 1992 with 1000
kg ha-1 of methyl bromide, using the procedure
described by Muchow et al. (1993). Cultivar Ql17
was planted on 23 July 1992. The plant (P) crop was
burnt and the site mechanically harvested on 18
August 1993 with an area sampled for yield by
hand-harvesting on 16 August 1993. The 1st ratoon
(1R) crop was sampled at regular intervals until final
harvest on 23 August 1994. The soil conditions at
the commencement of the P and 1R crop are shown
in Table 1. The nutrient status of the soil (other than
for N) changed little from the P to the 1R crop.
At planting, the sets were treated with
methoxyethyl mercury chloride at 0.1 kg ha -1 to
control fungi and planted with chlorpyrifos at 1.2 kg
ha ~ for control of wireworms. Row spacing was
1.48 m and plots were 40 m long by 8 rows for each
N rate with 2 replicates. Two replicates with the
sample sizes described below have been shown to
produce acceptable levels of precision (Muchow et
al., 1993). Plots were arranged in a 3 2 grid as a
randomised block design.
Fertiliser at planting was 56 kg N ha-~ as urea,
and 4 6 k g P h a -l , 8 4 k g K h a - 1 and 2 2 k g S h a 1
in a commercial planting formulation. For the P
crop, the three N rates were 56, 107 and 268 kg N
ha -~. In addition to the 56 kg N ha-1 at planting,
the 107 and 268 kg N ha ~ treatments received 51

145

R.C. Muchow et al. / Field Crops Research 47 (1996) 143-153

Table 1
Soil characteristics for 0-20 cm expressed as a mean of 6
locations sampled
Variate

Pre-plant 20 Post-harvest 6
July 1992
September 1993

pH (1:5 soil:water)
5.5
Electrical conductivity (dS m - 1) 0.04
Organic carbon (%)
0.8
NO 3 nitrogen (mg kg- J )
3.6
SO4 sulphur(mg kg- ] )
5
BSES P (mg k g - t)
92
Exchangeable Mg (me 100 g-] )
1.46
Exchangeable Ca (me 100 g t )
4.82
Exchangeable Na (me 100 g-] )
0.11
Exchangeable K (me 100 g J )
0.13
Exchangeable A1 (me 100 g - J )
0.06
CEC (me 100 g - i )
6.58

5.9
0.06
0.8
0.5
9
84
1.63
5.32
0.36
0.13
0.01
7.45

Determination methodology given in Kerr and Von Stieglitz

(1938).

kg N ha-~ as ammonium sulphate on 13 November

1992 and the 268 kg N ha-~ treatment received 161
kg N ha -j as urea on 23 September 1992. Boron
was applied (as borax) aerially from a boomspray at
1.5 kg ha -1 on 12 October 1992. On 12 October and
5 November, oxycarboxin at 375 g ha-~ was applied
for control of leaf rust. On 21 September, MCPA
amine was applied at 250 g ha-J to control vine
weeds. All plots began to lodge on 3 February 1993.
The 1R crop had three N treatments: 0, 55 and
774 kg N ha 1, which were superimposed upon the
low, medium and high N rate plots from the P crop.
The medium and high N rate in the 1R received 55
kg N ha J as urea immediately after harvest of the P
crops and the high rate received an additional 167

and 136 kg N ha i as urea on 21 October and 8

November 1993, respectively. The high N rate treatment also received 416 kg N ha 1 applied via the
trickle irrigation system (fertigated) as ammonium
nitrate on seven occasions (30 December 1993, 28
January, 9 March, 5 April, 10 May, 9 June and 27
July 1994 at ca. 60 kg N ha-l). The aim of the
fertigated high N treatment was to maintain a constant high level of soil mineral N supply throughout
the season, as soil mineral N in the high N rate in the
P crop was reduced to a similar level as the low and
medium N rates in the latter half of the season
(Table 3). In addition to fertiliser N applied, all plots
in the 1R crop received 131 kg K ha -1 and 53 kg S
ha-] as potassium sulphate on 20 September 1993.
On 16 November and 8, 10 and 16 December 1993,
oxycarboxin at 375 g ha-~ was applied for control
of leaf rust. On 16 December MCPA amine was
applied at 250 g ha -1 to control vine weeds. The
high N plots lodged in mid-June 1994, while the low
and medium N plots remained erect throughout the
season.
In the absence of rain, the crop in both seasons
was irrigated with overhead sprinklers with approximately 35 mm per week until November, and then
trickle irrigated with approximately 40 mm every
week (approximately equivalent to evapotranspiration) until harvest, so that soil water was not limiting
to growth.
2.2. Measurements

The P crop was sampled on 16 August 1993 (389

days after planting). The ratoon crop was sampled on

Table 2
Sampling dates for the 1R crop
Sampling date

Days after
ratooning

Mean solar radiation

(MJ m -2 d ])

Mean temperature
(C)

18/01/94
01/03/94
11/04/94
04/05/94
14/06/94
19/(/7/94
23/08/94

153
195
236
259
300
335
370

21.8
18.0
19.6
16.3
13.3
14.0
15.4

24.9
27.3
25.8
22.8
21.4
18.9
18.1

The mean daily climatic conditions from ratooning on 18 Aug. 1993 to the first sampling on 18 Jan. 1994, and for subsequent sampling
intervals are also given.

146

R.C. Muchow et al. / Field Crops Research 47 (1996) 143-153

seven occasions, as shown in Table 2. The sampling

procedure to measure yield accumulation was based
on that developed by Muchow et al. (1993), which
sought to balance data precision with manpower
requirements.
At each sampling, four inner rows each 2.5 m in
length were cut at ground level from each plot. Load
cells were used to determine the total fresh weight
(_+0.1 kg) in the field. A 15-stalk subsample, representative on a fresh weight basis, was taken and
partitioned into green leaf blades, dead leaf and dead
sheaths (defined as trash), millable stalks and cabbage (defined as the immature top of the stalk plus
green leaf sheaths). The fresh weight of each component was determined. Fresh millable stalk (cane)
yield was calculated as the product of the field
quadrat total fresh weight and the proportion of
millable stalks on a fresh weight basis. Then the
material from the leaf blades, cabbage and millable
stalk components was fibrated (finely-chopped) using a cutter-grinder, and two representative subsamples were placed in 850 ml aluminium foil trays for
drying at 80C. After the dry matter content was
determined, the green leaf, trash, millable stalk and
cabbage dry matter yield were determined per unit
land area from the total fresh weight, component
proportion of the total aboveground material on a
fresh weight basis, and the component dry matter
content. The net aboveground biomass was calculated as the sum of the individual components. Each
component was analysed for nitrogen concentration
using Kjeldahl digestion followed by ammonium
determination by the indophenol-blue colorimetric
procedure (Henzell et al., 1968). The net aboveground N accumulation was calculated as the sum of
the N accumulation in the individual plant components.
From the fresh fibrated material of millable stalk,
two further 500 g samples were taken for CCS
(Commercial Cane Sugar) and sucrose analysis. For
commercial sugarcane production in Australia, the
payment for sucrose is based on the fresh millable
stalk (cane) yield and CCS. The relationship between
CCS and the sucrose concentration in fresh millable
stalks is examined in this paper. The fibrated material was placed into a steel cylinder cage, and juice
was expressed by applying 15.7 MPa for 60 s using a
Carver Press (Model M, F.S. Carver, Wabash, IN,

USA). The press method was adopted in order to

standardise juice extraction and fibre determination
(Tanimoto, 1964; Walker, 1971). The juice was divided into three portions for determination of brix,
pol, and sucrose concentration using High Performance Liquid Chromatography (HPLC) following
the procedures set out by the International Committee for Uniform Methods of Sugar Analysis (1994).
The fresh and dry weight of the remaining fibrated
material (biscuit) was determined for calculation of
fibre content. Whilst this method differs from the
method of fibre determination used at Australian
sugar mills, it provides a standardised rapid procedure for fibre determination which gives similar
results to the Industry procedure (Walker, 1971).
Brix was determined on the juice using an automatic
temperature compensated (20C) Brix meter (Model
PR-1, Atago Pty Ltd, Tokyo, Japan). For the determination of Pol, 2.5 g of lead acetate was thoroughly
mixed with the juice, the juice was filtered through a
Whatman No. 91 filter paper, and the filtrate was
passed through a Polarimeter (Model AA5, Optical
Activity, Huntingdon, UK). For determination of
CCS, the following calculations were made:
F=

(100 DF) - (FF X B)

CCS =

5 X ( 100 - B)
(39XPX(95-F)
)
99.82 + (-0.-~-5XB)
- ( ( 0 . 5 X (B + ((0.00137

-(3E-05

X B ) ) X T 2)

+ ((0.00172 X B - 0.0044) X T)
- ( 0 . 0 2 2 4 X B) - 0 . 4 6 ) X(97 - F ) ) ) )

X 100 -1
where F is fibre percentage; DF is measured dry
biscuit weight; FF is measured fresh biscuit weight;
P is measured Pol in first expressed juice; B is
measured Brix in first expressed juice; and T is
20C.
For HPLC analysis, the juice was syringed through
a Swinnex filter (Millipore, Milford, MA, USA)
using 0.8 Ixm filter paper, and placed in vials that
were frozen prior to determination. The sucrose concentration in the juice was determined using double

R.C. Muchow et al. / Field Crops Research 47 (1996) 143-153

injection HPLC, and the extractable sucrose concentration in the stalk (g sucrose g-1 FW) was calculated as:

"~

(a)

500 - FF
Sucrose concentration ---

30

147

,,~

25

.~

20

~,
/,..-x

1",

I AA

i
--

i
i
1992/93

--

1993/94

-~

500 S

where FF is measured biscuit fresh weight after

pressing the 500 g of fibrated material and S is the
sucrose concentration in juice. The 'extractable' sucrose concentration in the stalk was calculated to
allow direct comparison with stalk CCS which is a
measure of recovery of sucrose by milling. The
'total' sucrose concentration was not calculated, but
may be calculated by substituting DF for FF in the
above formula, with the additional assumption that
the juice in the fresh biscuit weight has the same
sucrose concentration as the expressed juice with
pressing.
In the P crop, soil mineral N was sampled at 3
days before planting, and then 95, 223 and 317 days
after planting to a depth of 90 cm, which was the
approximate depth of the water table at this site. In
the 1R crop, soil mineral N was sampled at 16 days
after ratooning before any fertiliser was applied, and
then at 85, 246 and 414 day after ratooning. At each
sampling, six 45-mm diameter cores spanning the
space from one inter-row to the neighbouring interrow, were taken at two locations in each plot and
bulked. From the bulk sample, a subsample was
taken from the 0-20, 20-40, 4 0 - 6 0 and 60-90 cm
soil layers for extraction of mineral N with 1 M KCI
and a further sub-sample was taken for determination
of soil water content. The soil solution extracts were
analysed for nitrate and ammonium using the method
of Henzell et al. (1968) and converted to amounts of
mineral N per hectare to a depth of 90 cm by using
previously-determined bulk densities for the site.

3. R e s u l t s

3.1. Aerial and soil enuironment

The weekly climatic conditions measured at the

site for both crops are shown in Fig. 1. Mean
temperature and incident solar radiation were higher
during summer (December-February) in 1993-94
than in 1992-93; otherwise climatic conditions were

35
o

i
--

1992/93

--

1993/94

(b)

30

o 25
o

20
I

?
I

120011..... I
1000

(c

1992/93
[] 1 9 9 3 / 9 4

800

-~

600
400
200

..m

..~

I~

Ih

M A

...

M J

Fig. 1. W e e k l y means of daily solar radiation and mean temperature, and monthly totals of rainfall at Macknade for the 1992-93
and 1993-94 experiments.

similar for both crops. During winter, solar radiation

remained above 10 MJ m -2 day -1 and mean temperature remained above 15C. The climatic conditions between sampling intervals for the 1R crop are
shown in Table 2. The P crop had a growth duration
of 389 days and the average daily solar radiation was
18.6 MJ m -2 day -1 and mean temperature was
23.3C. The 1R crop had a growth duration of 370

Table 3
Mineral N ( 0 - 9 0 cm; kg ha -1 ) for Q117 planted at Macknade on
23 July 1992 and harvested on 16 August 1993
Days after

N applied (kg ha l )

planting

56

107

268

Significance

- 3
95
223
317

57.1
92.7
21.6
6.40

56.3
149
13.9
6.70

49.9
247
68.3
6.50

NS
NS
**
NS

The significance of the response to N applied, from analysis of

variance, is given where P < 0 . 0 5 * P < 0 . 0 1 **, P <
0.001 * * *, P > 0.05 NS.

R.C. Muchow et al. / Field Crops Research 47 (1996) 143-153

148

Table 4
Mineral N ( 0 - 9 0 cm; kg ha - 1 ) for Q117 ratooned at Macknade
on 18 August 1993 and harvested on 23 August 1994
Days after
ratooning
16
85
246
414

N applied (kg ha ~)
0
13.8
10.9
7.25
4.6

Significance

55

774

20.0
21.9
7.50
4.6

22.6
292
44.8
47.5

NS
*
***

The significance of the response to N applied, from analysis of

variance, is given where P < 0 . 0 5
*, P < 0 . 0 1 **, P <
0.001 * * *, P >_ 0.05 NS.

days and the average daily solar radiation was 18.5

MJ m -2 day -1 and the mean temperature was
23.5C. The total rainfall received for each crop
cycle was 1631 and 2007 mm in 1992-93 and
1993-94, respectively.
Prior to planting, the mean mineral N present in
the top 90 cm of the soil profile was 54.4 kg ha-1
(Table 3). By 95 days after planting (DAP), the
mineral N had increased in all treatments due to
fertilisation and mineralisation, despite some crop N
uptake. By 223 DAP, the mineral N was depleted in
all treatments, but it was significantly lower where
56 and 107 kg N ha -l was applied than where 268
kg ha-~ was applied. Very little mineral N remained
in the 0 - 9 0 cm soil profile at 317 DAP.
At ratooning and prior to fertilisation, the mineral
N level in the 0 - 9 0 cm soil profile was much lower
than at planting (Tables 3 and 4). Subsequent samplings after ratooning (Table 4) showed very little
mineral N present where 0 and 55 kg N ha-~ were
applied, but significantly more in the fertigated treatment. Interestingly, despite monthly fertigation, the
mineral N present at 246 d after ratooning (DAR)
was relatively low (44.8 kg N ha - l ) and remained
low until harvest (47.5 kg N ha -1 at 414 DAR)
despite the fertigated crop receiving a further five N
applications during late growth to ensure adequate
soil N availability for crop uptake.
3.2. Yield of P and 1R crop at final harvest

At harvest of the P crop at 389 DAP, there was a

significant increasing linear trend in cane yield with
increasing applied N and a decreasing linear trend in

CCS and stalk sucrose concentration (g g-1 FW)

(Table 5). However, there was no response in sucrose yield and crop biomass to N application. Furthermore, stalk sucrose concentration (g g - i DW)
was not significantly different at the three rates of
applied N (Table 5). Consequently, the decrease in
CCS and stalk sucrose concentration (g g - i FW)
with increasing N application was largely associated
with the significant decrease in stalk dry matter
content with increasing N application (Table 5).
Crop N uptake increased significantly with increasing N application resulting in an increase in stalk N
concentration (Table 5). Since sucrose yield and crop
biomass did not respond to N application, the efficiency of N-use, measured as the ratio of biomass to
crop N uptake (Biomass/N ratio) and as the ratio of
stalk sucrose yield to crop N uptake (Sucrose/N
ratio) decreased with increasing N application (Table
5).
The 1R crop responded differently to N application, with all variates except stalk sucrose concentration (g g - 1 DW) showing a significant response to N
application at the final harvest at 370 d after ratooning (Table 6). Both cane yield and sucrose yield
increased with increasing N application, despite a
depression in CCS and stalk sucrose concentration (g

Table 5
Crop response to nitrogen (N) applied for Q117 planted at Macknade on 23 July 1992 and harvested on 16 August 1993
Variate

N applied (kg h a - J )
56

107

Significance

268

Cane yield (t h a - l)
163
173
191
*
CCS (%)
16.1
14.9
13.1
Stalk sucrose (g g - 1 FW)
0.157 0.148 0 . 1 3 0 "
Stalk sucrose yield (t ha 1) 25.5
25.5
24.7 NS
0.517 0.484 NS
Stalk sucrose concentration
0.488
(g g - ~ DW)
Stalk dry matter content
0.322 0.286 0.268 *

(g DW g-~ FW)
Crop biomass(t ha- 1)
Crop N uptake (kg ha- 1)
Stalk N concentration
(mgNg-I DW)
Biomass/N ratio (t kg- J )
Sucrose/N ratio (t kg- i )

63.5
60.5
60.8 NS
140
190
256
**
1.400 2.200 3.300" *
0.454
0.183

0.327
0.146

0.238 * *
0.103 *

The significance of the linear trend of yield variate on N applied

is given where P < 0.05 *, P < 0.01 * *, P < 0.00l * * *, P >__
0.05 NS.

R.C. Muchow et al. / Field Crops Research 47 (1996) 143-153

Table 6
Crop response to nitrogen (N) applied for Q l 1 7 ratooned at
Macknade on 18 August 1993 and harvested on 23 August 1994
Variate

N applied (kg h a - 1)
0

Cane yield (t h a - 1)
CCS (%)
Stalk sucrose (g g - l FW)
Stalk sucrose yield (t h a - i )
Stalk sucrose concentration
(g g - t DW)
Stalk dry matter content
(g D W g ~ FW)
Crop biomass (t h a - 1 )
Crop N uptake (kg ha 1)
Stalk N concentration
(mg N g - l DW)
B i o m a s s / N ratio (t k g - 1)
S u c r o s e / N ratio (t k g - ~)

55

20

30.6
48.7
0.950

0.287 *

0.785
0.301

0.223 *
0.078 *

'

'

10

15

20

Fig. 3. Relationship between CCS and stalk sucrose concentration

(% FW). The solid 1 : 1 line is shown. The linear relationship fitted
to the data is: y = 1 . 2 6 3 ( _ + 0 . 0 4 0 ) x - 2 . 9 1 2 ( + 0 . 5 0 7 ) r 2 = 0.981.

The time-course of yield accumulation in response to N supply was measured for the 1R crop.
At this location, the commercial harvest season

.trA, I

ha-'

Stalk sucrose concentration (~ FW)

3.3. Time-course o f yield accumulation

200

,jkA.,///I/A

48.6
62.1
**
62.1
278
**
0.800
3.352 *

0.627
0.233

ha -~

10

0.333

kg

55 g s N

**
**
*
***
NS

g-J FW) in the fertigated treatment. Again this

depression was associated with a decrease in stalk
dry matter content. Crop N uptake and stalk N
concentration were highest in the fertigated treatment
with consequent lowest efficiency of N-use.

ranges from June until November. In this study,

samples were taken from January to August to assess
the impact of N on early sucrose accumulation.
Commercial yield in Australia is based on the weight
of fresh millable stalks (cane yield) and CCS which
is a measure of cane quality. Cane yield increased
with N-application at all samplings in this study (Fig.
2a). However, CCS was lowest at the highest N
application at all samplings (Fig. 2b). There was
little difference in CCS in the 0 and 55 kg N ha-1
treatments, except in January and March when it was
significantly higher in the 0 kg N h a - l treatment
(Fig. 2b). In these two treatments, CCS was above
10% in March, was 14.3% in April, and was above
17% in July-August.
Fig. 3 shows how CCS, which is calculated from
pol, brix and fibre, is related to actual stalk sucrose

20

(a)

774

Significance

The significance of the response to N applied from analysis of

variance is given where P < 0 . 0 5 *, P < 0 . 0 1
**, P <
0.001 *** P > _ 0 . 0 5 NS.

.~.

774

73
118
167
17.3
17.8
13.3
0.155
0.158
0.131
11.4
18.6
21.8
0.469
0.475
0.457
0.330

149

O0
I

kg

(b)

150

15

100

10

5o

@0

kg N ha -~

h a -1

Fig. 2. Accumulation of (a) cane yield and (b) CCS for Q117 ratooned at Macknade on 18 August 1993 and grown at three rates of applied
N. Error bars are twice the standard error of the mean where larger than the symbol.

R.C. Muchow et al. / Field Crops Research 47 (1996) 143-153

150
25

20

774 kg N ha "~
55 ks N ha -t

"00

plings, sucrose yield was highest at the highest rate

of applied N.
Biologically, yield accumulation in millable stalks
can be examined on a dry weight basis in terms of
stalk biomass and stalk sucrose concentration (g g DW). The response of stalk biomass to applied N
(Fig. 5a) was similar to the response of stalk sucrose
(Fig. 4). Stalk biomass was always lowest where no
N was applied. Similarly to stalk sucrose yield, only
after May was stalk biomass highest at the highest
rate of applied N.
The stalk sucrose concentration on a dry weight
basis increased in all N treatments until April; thereafter stalk sucrose concentration was relatively constant (Fig. 5b). At the early samplings, stalk sucrose
concentration (g g-~ DW) decreased with increasing
N application, but after April the tendency remained
but the differences were relatively small (Fig. 5b).
The fact that the pattern of accumulation of stalk
sucrose (Fig. 4) was similar to the pattern of accumulation of stalk biomass (Fig. 5a) in response to N
supply, indicates the dominant effect of biomass
accumulation rather than the partitioning of biomass
to sucrose (i.e., stalk sucrose concentration) in determining stalk sucrose accumulation.
The effect of N supply on stalk dry matter content
can help to explain the contrasting time-courses of
CCS and stalk sucrose concentration (g g ~ DW).
Stalk dry matter content was lowest in the highest N
application treatment at all samplings (Fig. 6). There
was little difference in stalk dry matter content in the
0 and 55 kg N ha -1 treatments. Interestingly, the
time-course in stalk dry matter content (Fig. 6) was

kg N ha-'

5
0

Fig. 4. Stalk sucrose a c c u m u l a t i o n for Q l 1 7 ratooned at M a c k nade on 18 A u g u s t 1993 a n d g r o w n at three rates o f applied N.
Error bars are twice the standard error o f the m e a n where larger
than the symbol.

concentration on a fresh weight basis, measured

using HPLC. Nitrogen supply had no effect on the
relationship which showed that there was relatively
close agreement between the measures in the range
of CCS values between 10% and 15%. At higher
values, CCS tended to overestimate sucrose concentration (Fig. 3 and Tables 5 and 6), and the converse
was true when CCS was below 10% (Fig. 3).
The effect of N supply on the time-course for
accumulation of millable stalk sucrose is shown in
Fig. 4. Where no N was applied, sucrose yield was
lowest at all samplings. However, there was no
difference in sucrose yield between the two treatments where N was applied up to May, when the
average yield was 15.3 t ha -~ . In effect, the higher
CCS compensated for the lower cane yield where 55
kg N ha-1 was applied. Thereafter, with later sam-

60

0.55

(b)

(a)
50

0.50

40

=~ 0 . 4 5

30

0.40

.~ 20

u 0.35

C"

=o
o

10

_ / w

~= 0.30

' t 5 5 k g N h a -1
0 kg N ha -~

'

'

'

0.25

/
,=/
m |

55

kg N h a -~
0 k g iN h a -~

Fig. 5. A c c u m u l a t i o n of (a) stalk b i o m a s s and (b) stalk sucrose concentration for Q 1 1 7 ratooned at M a c k n a d e on 18 A u g u s t 1993 and g r o w n
at three rates o f applied N. Error bars are twice the standard error o f the m e a n where larger than the symbol.

R. C. Muchow et al. / Field Crops Research 47 (1996) 143-153

0.35

m m i

and stalk biomass, and approaches the maximum

value of 0.49 g sucrose g-~ stalk biomass. The
relationship between stalk sucrose yield and cane
yield (Fig. 7b) was much more variable than the
relationship with stalk biomass (Fig. 7a). Stalk dry
matter content varies with crop age and N treatment
(Fig. 6) contributing to variation in the relationship
in Fig. 7b.

0.30

0.25

0.20

'~ 0.15
J

~m

m |

m i

4. Discussion

Fig. 6. Time-course of stalk dry matter content for Q117 ratooned

at Macknade on 18 August 1993 and grown at three rates of
applied N. Error bars are twice the standard error of the mean
where larger than the symbol.

This study has shown that N can be used to

manipulate the time-course of sucrose concentration
in fresh millable stalks or CCS during growth. Low
N supply increases CCS and this can be particularly
important where early harvest is contemplated. There
is clearly a trade-off in determining stalk sucrose
yield as low N supply also decreases fresh millable
stalk or cane yield. This study has shown that modest
N application can maximise both CCS and cane
yield at early harvest in May. Here, the 55 kg N
ha-~ treatment yielded 15.8 t h a - I of sucrose at a
CCS of 14.5% (Figs. 4 and 2b). However, with later
harvest in this study, higher N application significantly increased stalk sucrose yield albeit at lower
CCS. Maximising profitability in sugarcane production where a premium is paid for CCS, needs to
carefully consider the effect of N supply on the
time-course of accumulation of cane yield and CCS.
The effect of N supply on the time-course of stalk

similar to the time-course in CCS (Fig. 2b) in response to N supply.

3.4. Relationship between stalk sucrose yield and
stalk biomass and cane yield

The importance of stalk biomass accumulation in

determining stalk sucrose yield can be further examined by pooling the data from all samplings and N
treatments. The linear relationship in Fig. 7a shows
that 97% of the variation in stalk sucrose yield can
be accounted for by variation in stalk biomass due to
crop age and N treatment. The significant intercept
in the linear relationship indicates that stalk sucrose
concentration (g g-1 DW) increases with crop age

Ira)

774 kg N ha-'

(b) 774 kg N h a ~

l
~~

20

-~

10

.-~ 10

0
I0

20

30

40

50

S t a l k b i o m a s s (t ha ~)

60

../

Ass kg N ha-'

20

151

1/"',
l

50

I00

150

200

C a n e yield (t ha -~)

Fig. 7. Relationship between (a) stalk sucrose and stalk biomass and (b) stalk sucrose and cane yield (b) for Q117 ratooned at Macknade on
18 August 1993 and grown at three rates of applied N. The linear relationships fitted to the data are: (a) y ~ 0 . 4 8 9 ( _ + 0 . 0 1 4 ) x 1.167(_+0.408) r 2 = 0.970, (b) y = 0 . 1 4 0 ( _ + 0 . 0 1 2 ) x - 1.268(_+ 1.288) r 2 = 0.761.

152

R.C. Muchow et al. / Field Crops Research 47 (1996) 143-153

sucrose accumulation can largely be explained by the

effects of N on stalk biomass (Figs. 4 and 5aFig. 7a).
The production of biomass in sugarcane can be
examined in terms of the capture and utilisation of
solar radiation (Muchow et al., 1994), and low N
supply decreased both the capture and utilisation of
radiation in this study (Muchow et al., unpublished
data). Whilst increasing N supply did decrease the
sucrose concentration in dry millable stalks (Fig. 5b),
this effect was relatively small compared to the
dominant effect of N supply on stalk biomass. Consequently, during the time-course of yield accumulation, variation in stalk biomass accounted for most of
the variation in stalk sucrose yield irrespective of N
supply (Fig. 7a). The function fitted in Fig. 7a can be
used to model sucrose accumulation in relation to
stalk biomass. Importantly, the relationship in Fig.
7a also shows that sucrose accumulation in stalks is
a continuous process as stalks grow and biomass
increases, rather than any trigger (such as low temperature or water deficit) being required to promote
sucrose accumulation in stalks.
This study has shown that stalk dry matter content
has a major impact on the time-course of accumulation of commercial cane yield and CCS in response
to N supply. Under low N supply, stalk dry matter
content in the 1R crop increased from 21% to 33%
from January to August (Fig. 6). Under high N
supply, the corresponding range was 16% to 29%
(Fig. 6). The higher CCS early in the season under
low N supply can largely be explained with higher
stalk dry matter content. Similarly, at the final harvest of the P crop (Table 5), the increase in cane
yield and decrease in CCS with increasing N supply
can be largely explained by variation in stalk dry
matter content as there was no significant response
of stalk sucrose yield to N supply in the P crop.
Other studies (e.g. Das, 1936; Stevensen et al., 1992)
have also shown decreasing stalk dry matter content
with increasing N supply, with little effect of N
supply on stalk sucrose concentration on a dry weight
basis. Clearly, N has a marked effect of stalk dry
matter content and hence a greater effect on the
commercial measures (cane and CCS) of sugarcane
yield than the biological measures (stalk biomass and
stalk sucrose yield) of crop performance.
There was no response in crop biomass and stalk
sucrose yield to N treatment in the P crop but a large

response in the 1R crop (Tables 5 and 6). The

amount of mineral N contained in the soil profile
was much larger for the P crop than for the 1R crop
(Tables 3 and 4). Catchpoole and Keating (1995)
observed a similar contrasting response in two successive seasons and explained their results in terms
of the depletion of mineral N and a rundown in the
readily mineralisable pool of soil organic matter.
Even when crop biomass and sucrose yield did not
respond to N application, crop N uptake increased
with N application. This has been referred to as
luxury consumption of N (Stanford, 1963). This
resulted in an increase in stalk N concentration at
high N supply, as has been found by other workers
(Stevensen et al., 1992; Catchpoole and Keating,
1995).
Given luxury N uptake, the efficiency of use of N
uptake in biomass and sucrose production decreased
with high N application. Gascho et al. (1986) observed a similar decrease in sucrose N ratio with
increasing N application, with values ranging from
0.099 to 0.043 t ha -1 In order to develop the
physiological basis for maximising both N-use efficiency and crop yield, the challenge is to estimate
the minimum N uptake for maximum yield. Stanford
and Ayres (1964) developed a relationship from
Hawaiian crops to estimate the minimum N uptake
(x, kg ha - I ) associated with 98% of maximum crop
biomass (y, t h a - l ) : y = - 1 5 . 9 + 0.544x. For the
63 t ha-1 biomass produced by the P crop where 56
kg N ha -1 was applied, the minimum N uptake
calculated with this function is 145 kg N ha -1,
similar to that observed (Table 5). This suggests for
these crops that b i o m a s s / N ratios less than 0.4
indicate luxury N uptake and values greater than 0.4
indicate N deficiency. In the 1R crop where 0 and 55
kg N ha-J were applied, the b i o m a s s / N ratio was
higher than the 0.4 value and biomass and sucrose
yield did respond to additional N application (Table
6). Whilst maximum yield was not attained, the most
efficient use of N occurred in the 1R crop where 55
kg N ha i was applied, with 18.6 t ha -1 of sucrose
being produced for a crop N uptake only 62.1 kg N
ha 1. Whilst other studies indicate that sugarcane
generally exhibits a low (25-40%) recovery of applied fertiliser N (Keating et al., 1993), these results
indicate that sugarcane can use the N taken up
efficiently in the production of sucrose. The chal-

R.C. Muchow et al./Field Crops Research 47 (1996) 143-153

lenge remains to develop N management strategies

to maximise commercial yield with the most efficient use of nitrogen.

Acknowledgements
We thank the following for their contribution to
this study: Mike Spillman and CSR Technical Field
Department staff for technical assistance in the field;
Leonie Baker for sucrose analyses; May Ling Goode
for soil mineral N analyses; Cunningham Laboratory
Analytical Services staff for the plant N analyses;
and Heidi Vogelsang for assistance with data processing and presentation. The study was conducted
in partnership with the Sugar Research and Development Corporation.
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