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International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4149
EFFECTS OF FLOODING ON SUGARCANE (Saccharum officinarum L.)
PHYSIOLOGY, MORPHOLOGY, AND SUCROSE YIELD
Arinta R. Puspitasari1, SetyonoY. Tyasmoro2, Agung Nugroho2, Sri Winarsih1,
Ida Wenefrida3, and Herry S. Utomo3
1Indonesian Sugar Research Institute
2Faculty of Agriculture University of Brawijaya
3Louisiana State University Agricultural Center
ABSTRACT
Sugarcane is an important commodity in the world used for sugar and bioenergy. Weather
phenomenon, such as La Nina, and prolonged rainy seasons have impacted the cane plant and
sucrose yield and in many cases delayed ripening. The effects of four flooding periods were
studied in a replicated factorial design using four sugarcane varieties. The growth of
aboveground root increased as the duration of flood extended. The largest aboveground root
weight was produced by variety PSJT 941 when exposed to 12 weeks of flood. Each variety
responded to the flood treatments slightly differently in an aerenchyma number and stomata
density. Both upper and lower leaf surface stomata density were slightly affected by flood. PSJT
941 maintained a similar upper-leaf-surface stomata density throughout the treatments, except
during the 6-week flood treatment. As the flood durations increased, the proline content in the
leaves increased. A dramatic increase in the proline production was found in variety BL reaching
25.8 µM, which was four times higher than the proline content in the non-flooded (control) at the
end of the 12-week flood period. Flood treatments significantly affected sugarcane yield. Variety
PSJT 941 appeared very sensitive to the flood treatments. With 3 weeks of flood treatment, its
sugarcane yield reduced by 38% (3.55 kg). The sucrose yield of PSJT 941 also reduced
immediately just after 3 weeks of treatment, then further declined to 0.16 and 0.19 kg after 9 and
12 weeks of flood treatments, respectively. Dramatic effects of flooding were also found in
variety BL. Significant reduction of sucrose yield occurred just after 3 weeks of flooding and
continued to drop as the flood periods prolonged. After 12 weeks of flooding, its sucrose yield
was 0.03 kg (92% reduction). Varieties used in this study demonstrated differences in their
capabilities to respond to flooding. Even though there were no obvious physiological and
morphological traits that could directly be used as selection tools to breed for more flood-tolerant
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4150
varieties, the information obtained can be used to develop flood mitigation strategies for
sugarcane.
Keywords: sugarcane, aboveground root, stomata, aerenchyma, proline.
INTRODUCTION
Sugarcane is an important commodity in the world used for sugar and bioenergy. It is one of the
major C4 crops grown mainly in the tropic and subtropic regions. Main sugarcane production
countries in the world include Brazil with cane production of 34.1 million Mg of cane (34.1%),
India (15.8%), China (5.8%), Thailand (4.6), Pakistan (2.9%), Mexico (2.8%), Colombia (1.6%),
Indonesia (1.6%), Philippines (1.5%), and USA (1.3%) (FAO, 2013; Factfish, 2015). Due to
urbanization and land use change, sugarcane planting areas in many tropical regions have shifted
to less arable dry lands causing dependency on rainfall for successful production. Weather and
climate related events, such as a wider range of extreme temperature, precipitation, and other
extreme weather, are the key factors for sugarcane production, especially in many developing
countries. Weather patterns, such as the La Nina phenomenon or prolonged rainy season, give
unfavorable impacts to sugarcane and sucrose yield and in many cases delays sugarcane
ripening.
On land with a bad drainage system, high rainfall often causes flooding. The degree of plant
damage associated with flooding is determined by many factors, including the depth and duration
of flooding, flow of water in the soil, and changes in the physical, chemical, and biological
structure of soil (Tetsushi and Karim, 2007; Parent et al., 2008; Ren et al., 2014). Prolonged
flooding can affect sugarcane yield and its components through changes in the plant anatomy,
physiology, and metabolism. The levels of damage also depend on the growth phase of the plant.
In recent studies, flooding has caused a high rate of stem mortality, low growth rate, and reduced
yield (Islam et al., 2011a). Sugarcane under flooded conditions experiences significant changes
in roots morphology, such as an increase in fibrous root growth. Flooding conditions also induce
the formation and modification of root aerenchyma, an important spongy tissue that form spaces
or air channels that allows gas exchanges and root activity to sustain under flooded land
(Webster and Eavis, 1972; and Begum et al., 2013). The channels of air-filled cavities provide a
low-resistance internal pathway for the exchange of gases, such as oxygen and ethylene, between
the plant above the water and the submerged tissues. Aerenchyma is widespread in aquatic and
wetland plants which must grow in hypoxic soils (Keddy, 2010; Kozlowski and Pallardy, 1984).
Flooding decreases the rates of transpiration due to the closing of stomata, reduces
photosynthesis rates due to the decrease in effective leaf surface area, decreases plant growth
rate, and increases respiratory rates especially in the plant’s organs that are flooded. A
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4151
modification in respiratory metabolism from aerobe to anaerobe is one of the main impacts of
oxygen deficiency associated with flooding. In addition, flooding can cause an increase in the
percentage of coir and content instead of sugar in the sugarcane stem (Islam et al., 2011a,b).
Flooding can cause essential nutrients, such as nitrogen, phosphorus, and potassium, to leach.
Nitrogen deficiency as a result of leaching becomes the limiting factors in sugarcane production
under flooded conditions (Wiedenfeld and Enciso, 2008). Although stagnant water has no
influence on phosphorus and potassium content in leaves and stem (Gomathi et al., 2010), it
causes a significant reduction in nitrogen content in both leaves and stems by 28.07% and
29.53%, respectively. Prolonged stagnant water can also damage the respiratory roots by
causing the formation of toxic compounds that hamper the nutrient uptake. The objectives of this
study were to 1) evaluate the effects of flooding on several sugarcane varieties and 2) determine
physiological responses affected by flooding by specifically measuring stomata density, proline
content, and the structure of aerenchyma tissue.
MATERIALS AND METHODS
Physiological effects of continuous flooding on a sugarcane plant were studied in the greenhouse
in June 2015 - June 2016 at the Center for Sugarcane Plantation Research of Indonesia (P3GI)
near Pasuruan, Indonesia. The studies were arranged in a factorial complete randomized block
design with 2 factors (i.e. variety and duration of flooding) and 3 replications. Four sugarcane
varieties used in the studies were PS 881, PS 851, BL, and PSJT 941. The second factor was
duration of flooding 0, 3, 6, 9, and 12 weeks. Materials used in this study include sugarcane seed,
growing media (consisted of a mixture of soil, sand, and compost with ratio 1:1:1), and 50-liter
pots (47 cm in height and 41 cm in diameter) equipped with a faucet. Tools used were a
spectrophotometer, microscope, and equipment to measure brix, pol, and sugar levels. Flood
treatments were applied when the plants reach the age of 4 months. The experimental units
consisted of 3 potted plants. Data were collected at the end of each treatment as specified in the
experimental design and analyzed according to the design model. If the F test results indicated a
significant difference among treatments, the least significant difference (LSD, p<0.05 level) tests
were carried out.
Physiological variables measured in these studies include aboveground root weight, stomata
density on both the upper and lower surface of leaves, aerenchyma tissue, proline content,
sugarcane yield, and sugar content. Aboveground root weight was measured at the end of each
flood treatment. Stomata density was calculated under a microscope with a magnification of
400x. Proline content was measured according the protocol developed by Bates et al. (1970).
Briefly, 0.2 g fresh leaf tissue was extracted into 2 ml of 3% sulphosalicylic acid followed by
centrifugation. The resulted aliquot was put into a test tube, and 2 ml of Ninhydrin was added
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4152
followed by 2 ml of glacial acetic acid. The test tube containing the mixture was then heated to
boiling for 1 hour. The tube was then placed in a tub of ice. Five ml of toluene was added and
homogenized. The solution was read using a spectrophotometer in 520 nm to determine their
proline content. Aerenchyma tissue was evaluated under a microscope with a magnification of
400x. Sugarcane yield was calculated from the weight of stem per clump. Sucrose yield was
calculated by multiplying the sugarcane yield by sugar content. Analysis of statistic used was the
LSD test in significance level of 5%.
RESULTS AND DISCUSSION
Results:
Aboveground Sugarcane Root
Roots in the sugarcane stem that are near the ground grew as a response to extended flooding.
The growth of the aboveground root increased as the duration of flooding prolonged (Table 1).
When flooded, sugarcane produced three types of adventitious roots.
Table 1: Aboveground root weight of four sugarcane (Saccharum officinarum L.) varieties
(BL, PS 851, PS 881, and PSJT 941) following 3, 6, 9, and 12 weeks of continuous flooding.
Variety
Root Weight (g)†
Flood Duration (week)
0 3 6 9 12
BL
0
a
11.66
bcd
18.71
d
30.71
ef
54.54
g
PS 851 0 a 8.99 b 13.61 bcd 29.72 e 51.28 g
PS 881 0 a 11.07 bc 17.85 cd 42.45 fg 75.52 h
PSJT 941 0 a 16.93 cd 30.39 e 69.60 h 142.99 i
BNT 5% 0.97
KK (%) 12.25
†Mean values with the same letter(s) in the same column and row indicate not significantly
different based on LSD test, p< 5%.
The first type of root grew from the nodes under the water just a few day following flooding,
white in color then changed to pink. The top node produced these roots the most both in length
and size. The bottom nodes produced less amount of roots. The second type of root grew from
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4153
the first type, typically numerous, small in size, thin, grew upward against gravity. The third type
of root grew under prolonged flooding emerging at the aerial nodes, at the 1st - 3rd nodes above
the water level. These roots were few in number, hard, and short (3–5 cm length). The
aboveground root weight among the four varieties was significantly different as they responded
to the flood treatments (Table 1). The aboveground root weight varied among the varieties tested,
ranging from 51.28 g to 142.99 g. The largest aboveground root weight was produced by variety
PSJT 941 (142.99 g) when exposed to 12 weeks of continuous flood. PS 851 had the lowest
aboveground root weight in the 12-week treatment (51.3 g), which was significantly lower than
PSJT 941 and PS 881. PS 881 produced an aboveground sugarcane root mass of 75.52 g.
Stomata Density
In the normal non-flooded conditions, the four varieties (BL, PS 581, PS 881, and PSJT 941) had
stomata densities that were different and specific to each variety (Table 2). PS 881 had the
densest stomata on its upper leaf surface (30.09 μm-2) compared to the three other varieties. Its
stomata density on the lower leaf surface was 60.17 µm-2, which was higher than that of BL and
PS 851, but similar to that of PSJT 941 (57.82 µm-2). The stomata density on the upper leaf
surface of BL and PS 851 was not affected by flood treatments (3, 6, 9, and 12 weeks). PS 881
did not show any changes in its upper leaf surface stomata density following the flood treatments
of up to 9 weeks. After 12 weeks of flood treatment, however, its stomata density reduced to
23.97 µm-2. PSJT 941 maintained a similar upper leaf surface stomata density throughout the
treatments, except during the 6-week flood treatment (27.78 µm-2).
The stomata density on the lower leaf surface of PS 851 were not affected by the 3, 6, and 9
weeks of flood treatments. Its stomata density however was lower following the 12-week flood
treatment (41.68 µm-2). While PS 881 responded differently to 3, 6, and 9 weeks of flood
treatment, its lower leaf surface stomata density remained the same as the untreated check when
it was exposed to 12 weeks of flood treatment (Table 2). PSJT 941 had a significant reduction in
its lower leaf surface stomata density following 3, 6, and 12 weeks of flood treatments.
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4154
Table 2: Stomata density on the upper and lower leaf surface of four sugarcane
(Saccharum officinarum L.) varieties (BL, PS 851, PS 881, and PSJT 941) following 3, 6, 9,
and 12 weeks of continuous flooding.
A. Upper leaf surface
Variety
Number of Stomata†
(µm-²)
Flood Duration (week)
0 3 6 9 12
BL 25.39 cde 24.76 cde 23.19 bcde 24.52 cde 22.88 abcd
PS 851 22.88 abcd 23.50 cde 19.74 ab 22.56 abcd 18.80 a
PS 881 30.09 fgh 31.03 fgh 32.44 h 29.62 fgh 23.97 cde
PSJT 941
22.56
abcd
21.78
abc
27.27
efg
26.33
def
24.13
ce
BNT 5% 4.15
KK
10.08
B. Lower leaf surface
Variety
Number of Stomata†
(µm-²)
Flood Duration (week)
0 3 6 9 12
BL 54.84 ghi 49.36 cdef 50.14 ef 55.63 hij 52.65 fgh
PS 851 47.01 bcde 44.19 ab 46.54 bcd 44.19 ab 41.68 a
PS 881 60.17 k 59.23 jk 71.92 l 50.77 ef 60.64 k
PSJT 941
57.82
ijk
45.76
bc
51.48
fg
55.00
ghi
43.25
ab
BNT 5% 3.97
KK
4.60
†Mean values with the same letter(s) in the same column and row indicate not significantly
different based on LSD test, p< 5%.
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4155
Aerenchyma Tissue in Sugarcane Root
In response to flooding, sugarcane plants produced adventitious roots with well-developed aerenchyma. The formation of aerenchyma
in the cortex of these roots is an adaptation to less favorable growth conditions by increasing their porosity under prolonged flooding.
Figures 1-4 show the cross-section of aerenchyma tissue of BL, PS 851, PS 881, and PSJT 941 from flood treatments of 0, 3, 6, 9, and
12 weeks.
Figure 1: Cross-section of aerenchyma tissue of PS 881 after flooding: a. 0 Week, b. 3 Weeks,
c. 6 Weeks, d. 9 Weeks, and e. 12 Weeks (400X Magnification).
a b c d e
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4156
Figure 2: Cross-section of aerenchyme tissue of PS 851 after flooding: a. 0 Week, b. 3 Weeks, c. 6 Weeks,
d. 9 Weeks, and e. 12 Weeks (400X Magnification).
a b c d e
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4157
Figure 3: Cross-section of aerenchyma tissue of BL after flooding: a. 0 Week, b. 3 Weeks, c. 6 Weeks,
d. 9 Weeks, and e. 12 Weeks (400X Magnification).
a b c d e
Figure 4: Cross-section of aerenchyma tissue of PSJT 941 after flooding: a. 0 Week, b. 3 Weeks, c. 6 Weeks,
d. 9 Weeks, and e. 12 Weeks (400X Magnification).
a b c d e
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
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Each variety responded to the flood treatments slightly differently in number of aerenchyma cells
(Table 3). Among the four varieties evaluated, PS 881 has the highest number of aerenchyma
(51.0 units) in the non-flooded setting. The number of aerenchyma in BL was not affected by
treatment of 3, 9, and 12 weeks of flooding. The number was higher than the control following
the 6-week flood treatment (44 units). When exposed to 6 and 12 weeks of flooding, PS 851 had
the same aerenchyma number as the untreated. After the 3-week flood treatment, its aerenchyma
number was lower than the control, but increased after the 9-week flood treatment. The
aerenchyma number in PS 881 was unaffected by the 3-week flood treatment. As the treatment
was getting longer, the aerenchyma number decreased; 31.5 units after 9 weeks of flooding and
33 units after 12 weeks of flooding. PJST 941 had its aerenchyma number un-affected
throughout the treatments, except in the 6-week flood treatment where the aerenchyma number
was 52.5 units, significantly higher than that of the untreated one.
Table 3: Number of aerenchyma constituent cavities of four sugarcane (Saccharum
officinarum L.) varieties (BL, PS 851, PS 881, and PSJT 941) following 3, 6, 9, and 12 weeks
of continuous flooding.
Variety
Number of Aerenchyma Constituent Cavities†
(unit)
Flood Duration (week)
0 3 6 9 12
BL 35.5 bcd 31.5 ab 44.0 efg 37.5 bcde 35.0 bcd
PS 851 37.5 bcde 25.0 a 45.0 efgh 46.0 fgh 40.0 cdef
PS 881 51.0 gh 50.5 gh 41.5 def 31.5 ab 33.0 bc
PSJT 941
41.5
def
44.0
efg
52.5
h
38.5
bcdef
40.5
cdef
BNT 5% 7.74
KK 11.30
†Mean values with the same letter(s) in the same column and row indicate not significantly
different based on LSD test, p< 5%.
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
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Proline Content
It is well established that proline metabolism leads to an increase of mitochondrial reactive
oxygen species (ROS) production. Proline metabolism impacts cell survival and cell death in
different species (Miller et al., 2009; Cecchini et al., 2011; Phang and Liu, 2012). Proline’s
protective effect during stress is especially well documented in plants (Aleksza et al., 2017;
Liang et al., 2013.). Results from this study indicated that proline contents in the leaves increased
under flood conditions by varying magnitudes among different varieties (Table 4, Figure 5).
Each variety responded differently (Figure 5). In general, proline content increased as the flood
duration increased. A dramatic increase in proline production was found in variety BL reaching
25.8 µM, which was four times higher than the proline content in non-flooded (control) at the
end of the 12-week flood period. PS 851 had a proline content of 20.3 µM, which was a 75%
increase, while PS 881 and PSJT 941 had a modest increase of 50% (15.6 µM) and 40% (24.6
µM), respectively.
Table 4: Proline content of four sugarcane (Saccharum officinarum L.) varieties (BL, PS
851, PS 881, and PSJT 941) following 3, 6, 9, and 12 weeks of continuous flooding.
Variety
Proline Content†
(µM)
Flood Duration (week)
0 3 6 9 12
BL
6.2
a
12.1
bc
13.3
cde
14.9
def
25.8
l
PS 851 13.4 cde 14.5 cde 17.3 fgh 18.9 hi 20.3 ij
PS 881 10.7 b 12.1 bc 12.1 bc 12.9 bcd 15.6 efg
PSJT 941
17.6
gh
21.9
Jk
23.2
kl
24.4
kl
24.6
l
BNT 5% 2.61
KK
9.50
†Mean values with the same letter(s) in the same column and row indicate not significantly
different based on LSD test, p< 5%.
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4160
Figure 5: Relationship between varieties and duration of flooding on proline content.
Sugarcane Yield
Flood treatments significantly affected the Sugarcane Yield (SCY) with varieties responding
differently to the flood treatments (Table 5). While 3 weeks of a continuous flood treatment did
not affect the SCY of variety BL, the 6-week treatment caused a significant reduction in its SCY
(2.3 kg). Its SCY values remained at the same levels for the longer durations of 9 and 12 weeks
of flood treatments. Both PS 851 and PS 881 had their SCY un-affected throughout the
treatments from 3, 6, 9, and 12 weeks of continuous flooding. After 12 weeks of flooding, PS
851 had a SCY of 4.24 kg and PS 881 of 3.55 kg (Table 5). Variety PSJT 941 appeared very
sensitive to the flood treatment. Its SCY was reduced by 38% (3.55 kg) with just 3 weeks of
flood treatment. However, its SCY stayed at the same levels with longer flood periods of
treatments (6, 9, and 12 weeks).
y = 1.398x + 6.0868R² = 0.8644
y = 0.6063x + 13.239R² = 0.9824
y = 0.3541x + 10.558R² = 0.8494
y = 0.5468x + 19.054R² = 0.83
0
2
4
6
8
10
12
14
16
18
20
22
24
26
28
0 3 6 9 12
Pro
line
(µ
M)
Duration of Flooding (Week)
BL PS 851 PS 881 PSJT 941
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
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Table 5: Sugarcane yield of four sugarcane (Saccharum officinarum L.) varieties (BL, PS
851, PS 881, and PSJT 941) following 3, 6, 9, and 12 weeks of continuous flooding.
Variety
Sugarcane Yield (SCY) †
(kg)
Flood Duration (week)
0 3 6 9 12
BL
3.91
cd
4.24
cde
2.30
ab
2.02
a
1.58
a
PS 851 4.45 cde 3.86 cd 5.28 ef 4.03 cd 4.24 cde
PS 881 3.90 cd 4.50 cde 4.51 cde 4.50 cde 3.55 c
PSJT 941
5.73
f
3.55
c
4.80
def
3.48
bc
3.63
cd
BNT 5%
1.19
KK (%)
18.41
†Mean values with the same letter(s) in the same column and row indicate not significantly
different based on LSD test, p< 5%.
Sucrose Yield
Under non-flooded conditions, PSJT 941 had the highest sucrose yield (SY) of 0.66 kg, which
was significantly higher than the three other varieties (Table 6). BL, PS 851, and PS 881 had
similar sugar yields; BL (0.38 kg), PS 851 (0.35 kg), and PS 881 (0.29 kg). The SY of variety
BL reduced significantly just after 3 weeks of flooding and continued to drop as the flooding
periods prolonged. After 12 weeks of flooding, its SY was only 0.03 kg (92% reduction). PS 851
also experienced a significant reduction in SY when exposed to flooding in just 3 weeks. With
longer flooding periods of 6, 9, and 12 weeks, however, PS 851 maintained its SY as it was after
3 weeks of flooding. The SY of PS 881 stayed relatively the same levels following 3, 6, and 9
weeks of flooding, but declined to 0.16 kg after 12 weeks. Just like PS 851, the SY of PSJT 941
reduced immediately just after the first flood treatment (3 weeks), then further declined to 0.16
and 0.19 kg after 9 and 12 weeks of flood treatments, respectively.
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
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Table 6: Sucrose yield of four sugarcane (Saccharum officinarum L.) varieties (BL, PS 851,
PS 881, and PSJT 941) following 3, 6, 9, and 12 weeks of continuous flooding.
Variety
Sucrose Yield (SY) †
(kg)
Duration of Flooding(week)
0 3 6 9 12
BL
0.38
i
0.27
efgh
0.14
bc
0.09
ab
0.03
a
PS 851 0.35 hi 0.22 cdefg 0.26 defgh 0.20 bcdefg 0.21 cdefg
PS 881 0.29 fghi 0.30 fghi 0.29 fghi 0.26 defgh 0.16 bcd
PSJT 941
0.66
j
0.30
ghi
0.34
hi
0.16
bcde
0.19
bcdef
BNT 5%
0.11
KK (%)
26.05
†Mean values with the same letter(s) in the same column and row indicate not significantly
different based on LSD test, p< 5%.
Discussion:
Sugarcane develops ways to cope with less favorable growing conditions. Under flood situations,
sugarcane produced aboveground roots in attempt to offset negative effects of anaerobic
conditions. Reduction in primary root weight and stimulating adventitious root are a common
response to the flood (Jaiphong et al., 2016; Gilbert et al., 2007). In this study, the amount of the
aboveground roots differed by variety. The biggest aboveground root mass was produced by
PSJT 941; the high yielding variety. The medium yielding varieties produced moderate levels of
aboveground root mass when exposed to prolonged flooding. Results from previous studies by
others (Gilbert et al., 2007; Srinivasan and Batcha, 1962; Webster and Eavis, 1972), showed that
long-term flooding reduced leaf area index (LAI) and leaf weight. Both upper and lower leaf
surface stomata densities were slightly affected by flood. PSJT 941 maintained a similar upper
leaf surface stomata density throughout the treatments, except during the 6-week flood treatment
(27.78 µm-2). While PS 881 responded differently to 3, 6, and 9 weeks of flooding with its lower
leaf surface stomata density remaining the same as the untreated check when it was exposed to
12 weeks of flooding.
The aerenchyma number in variety BL was not affected by 3, 9, and 12 weeks of flooding
treatments. However, the number increased following the 6-week flood treatment (44 units).
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
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PJST 941 had a similar pattern. PS 881, however, had less aerenchyma as flooding prolonged.
Glaz and Gilbert (2006) and Glaz et al. (2004a,b) indicated that constitutive formation of stalk
aerenchyma could be an important adaptation that enables sugarcane to tolerate periodic floods.
Despite the presence of stalk aerenchyma 50-75% up the stalk, neither cultivar studied was able
to maintain yields when subjected to a 3-month summer flood (Gilbert et al., 2007). The results
reported, however, were slightly different from our findings. Long-term flooding caused
prolonged anoxia in the root zone leading to stress levels that were hard to overcome.
Proline accumulation in plants under various stresses is positively correlated with oxidative
stress tolerance (Anjum et al., 2012; Theocharis et al., 2012; Xu et al., 2012; Saeedipour, 2013).
Proline also plays a role in the post-stress recovery process. In general, proline content increased
as the flood durations increased. PS 851 had a proline content of 20.3 µM, which was a 75%
increase, while PS 881 and PSJT 941 had a modest increase of 50% (15.6 µM) and 40% (24.6
µM), respectively. A dramatic increase in proline production was found in variety BL reaching
25.8 µM, which was four times higher than the proline content in non-flooded (control) at the
end of the 12-week flood period. The four fold increase in proline accumulation found in the
least flood-tolerant variety BL could be a sign of tremendous stress to overcome.
Prolonged inundation, especially if the sugarcane is in the early stages of growth, can have
devastating consequences. Both PS 851 and PS 881 had their SCY un-affected throughout the
treatments from 3, 6, 9, and 12 weeks of continuous flooding. After 12 weeks of flooding,
however, PS 851 had a SCY of 4.24 kg and PS 881 of 3.55 kg. Variety PSJT 941 appeared very
sensitive to the flood treatment. Its SCY was reduced by 38% (3.55 kg) within just 3 weeks of
flooding. With longer flood periods of treatments (6, 9, and 12 weeks), however, its SCY
remained at the same levels. A previous field study by Gilbert et al. (2008) indicated that
flooding sugarcane in the summer caused sequentially greater yield reductions throughout the
harvest season in planted cane. Other reports (BSES, Information Sheet IS13015) stated that
cane may suffer around 15-20% yield loss after 5 days of submergence, between 30 and 60%
yield loss after 10 days, and between 37-100% yield loss after 15 days. The magnitude of loss for
each period of inundation depends on stalk height with the least loss for 2.5 m stalks and the
most loss for 0.5 m stalks for each period of inundation.
The sucrose yield (SY) of PSJT 941 reduced immediately just after the first flood treatment (3
weeks), then further declined to 0.16 and 0.19 kg after 9 and 12 weeks of flood treatments,
respectively. Dramatic effects of continuous flooding was also found in variety BL. Significant
reduction of SY occurred just after 3 weeks of flooding, and continued to drop as the flood
periods prolonged. After 12 weeks of flooding, its SY dropped to 0.03 kg (92% reduction). As
comparisons, field studies conducted by Gilbert et al. (2007) showed that SY for flooded cane,
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
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compared with the non-flooded control, were 9.6 t sucrose ha-1 versus 11.7 t sucrose ha-1 early,
9.2 t sucrose ha-1 versus 12.8 t sucrose ha-1 mid-season, and 7.8 t sucrose ha-1 versus 12.3 t
sucrose ha-1 at late harvest. The flooding sugarcane in the summer caused sequentially greater
yield reductions throughout the harvest season in planted cane.
One of the most effective ways to address the prolonged flooding caused by more frequent
abnormal climate patterns is to develop new flood-tolerant cultivars. Varieties used in this study
demonstrated their differences in the capability to respond to flooding. Introgression and
accumulation of various genes contributing to flood tolerance is the key in developing new
flood-tolerant cultivars. Even though there are no obvious physiological and morphological traits
that can directly aid in selecting and breeding for more flood-tolerant varieties, the information
obtained from this study can be used as a device to develop effective flood mitigation strategies
for sugarcane.
REFERENCES
Aleksza, D., Horváth, G.V., Sándor, G., and Szabados, L. 2017. Proline accumulation is
regulated by transcription factors associated with phosphate starvation. Plant Physiol.
175(1): 555–567.
Anjum, S.A., Farooq, M., Xie, X.Y., Lie, X.J., and Ijaz, M.F. 2012. Antioxidant defense system
and proline accumulation enables hot pepper to perform better under drought. Sci. Hort.
140:66-73.
Bates, L.S., Waldren, R.P., and Teare, I.D. 1973. Rapid determination of free proline for water-
stress studies. Plant Soil (1973) 39:205.
Begum, M.K., Alam, M.R. and Islam, M.S. 2013. Adaptive mechanism of sugarcane genotypes
under flood stress condition. World J. of Agri. Sci., 1(2):56-64.
BSES, Information Sheet IS13015. Managing flood damage cane.
http://www.canegrowers.com.au/icms_docs/147481_BSES_Managing_flood_damaged_ca
ne.pdf
Cecchini, N.M., Monteoliva, M.I., and Alvarez, M.E. 2011. Proline dehydrogenase is a positive
regulator of cell death in different kingdoms. Plant Signal Behav. 6:1195-1197.
Factfish. 2015. “Sugarcane, production quantity (tons) - for all countries”.
http://www.factfish.com/statistic/sugar+cane,+production+quantity.
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4165
FAO, “Report, Food and Agricultural Organization, United Nations: Economic and Social
Department,” The Statistical Division, FAO 2013 FAOSTAT, 2014, http://faostat3.fao.org/
home/index.html#DOWNLOAD.
Gilbert, R.A., Rainbolt, C.R., Morris, D.R., and Bennett, A.C. 2007. Morphological responses of
sugarcane to long-term flooding. Agron. J. 99:1622-1628.
Gilbert, R.A., Rainbolt, C.R., Morris, D.R. and McCray, J.M. 2008. Sugarcane growth and yield
responses to a 3-month summer flood. Agricultural Water Management 95:283-291.
Glaz, B. and Gilbert, R.A. 2006. Sugarcane response to water table, periodic flood, and foliar
nitrogen on organic soil. Agron. J. 98:616-621.
Glaz, B., Morris, D.R., and Daroub, S.H. 2004a. Periodic flooding and water table effects on two
sugarcane genotypes. Agron. J. 96:832-838.
Glaz, B., Morris, D.R., and Daroub, S.H. 2004b. Sugarcane photosynthesis, transpiration and
stomatal conductance due to flooding and water table. Crop Sci. 44:1633-1641.
Gomathi, R., Chandran, K., Rao, P.N.G. and Rakkiyappan, P. 2010. Effect of waterlogging in
sugarcane and its management. Sugarcane Breeding Institute. Coimbatore. 4 p.
Jaiphong, T., Tominaga, J., Watanabe, K. Nakabaru, M., Takaragawa, H., Suwa, R., Uenoa, M.
and Kawamitsu, Y. 2016. Effects of duration and combination of drought and flood
conditions on leaf photosynthesis, growth and sugar content in sugarcane. Plant Production
Science 19(3):427–437.
Islam, M.S., Miah, M.A.S., Begum, M.K., Alam, M.R. and Arefin, M.S. 2011a. Growth, yield
and juice quality of some selected sugarcane clones under waterlogging stress condition.
World J. or Agric. Sci., 7(4):504-509.
Islam, M.S., Miah, M.A.S., Begum, M.K., Alam, M.R. and Arefin, M.S. 2011b. Biochemical
studies of juices quality and yield performance of some promising sugarcane clones under
water-logging stress condition. J. Agro. Environ., 5(1):87-90.
Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge
University Press, Cambridge, UK. 497 pp.
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4166
Kozlowski, T.T. and Pallardy, S.G. (1984). Effects of flooding on water, carbohydrate and
mineral relations. In Flooding and plant growth. T.T. Kozlowski (ed). pp. 165–193.
Academic Press Inc., Orlando, Florida.
Liang, X., L. Zhang, S.K. Natarajan, and Becker, D.F. 2013. Proline mechanisms of stress
survival. Antioxid Redox Signal 19(9): 998–1011.
Miller, G., Honig, A., Stein, H., Suzuki, N., Mittler, R., and Zilberstein, A. 2009. Unraveling
delta1-pyrroline-5-carboxylate-proline cycle in plants by uncoupled expression of proline
oxidation enzymes. J. Biol. Chem. 284:26482-26492.
Parent, C., N. Capelli, A. Berger, M. Crevecouer, and Dat, J.F. 2008. An overview of plant
responses to soil waterlodging. Plant Stress 2(1):20-27.
Phang, J.M., Liu, W., Hancock, C., and Christian, K.J. 2012. The proline regulatory axis and
cancer. Front Oncol. 2:60.
Ren, B., Zhang, J., Li, X., Fan, X., Dong, S., Liu, P., and Zhao, B. 2014. Effects of waterlogging
on the yield and growth of summer maize under field conditions. Can. J. Plant Sci. 94:2331.
Saeedipour, S. 2013. Relationship of grain yield, ABA and proline accumulation in tolerant and
sensitive wheat cultivars as affected by water stress. PNAS India. 10.1007/s40011-012-
0147-5.
Srinivasan, K. and Batcha, M.B.G.R. 1962. Performance of clones of Saccharum species and
allied genera under conditions of water-logging. Proc. Int. Soc. Sugar Cane Technol. 11:
571-577.
Tetsushi, H. and Karim, A. 2007. Flooding Tolerance of Sugarcane in Relation on Growth,
Physiology and Root Structure. South Pacific Studies, 28(1):9-22.
Theocharis, A., Clement, C., and Barka, E.A. 2012. Physiological and molecular changes in
plants grown at low temperature. Planta 235:1091-1105.
Wiedenfeld, B. and Enciso, J. 2008. Sugarcane responses to irrigation and nitrogen in semiarid
south Texas. Agronomy Journal, v.100, p. 665-671.
Webster, P.W.D. and Eavis, B.W. 1972. Effects of flooding on sugarcane growth. I. Stage of
growth and duration of flooding. Proc. Int. Soc. Sugar Cane Technol. 14:708-714.
International Journal of Agriculture and Environmental Research
ISSN: 2455-6939
Volume:03, Issue:06 "November-December 2017"
www.ijaer.in Copyright © IJAER 2017, All right reserved Page 4167
Xu, F.Y., Wang, X.L., Wu, Q.X., Zhang, X.R., and Wang, L.H. 2012. Physiological responses
differences of different genotype sesames to flooding stress. Adv. J. Food Sci. Technol.
4:352-356.