Documente Academic
Documente Profesional
Documente Cultură
FLOODING STRESS
Keywords
plant senescence; planting pattern; root
oxidizing activity; single seedling; system of
rice intensification; water regime
Correspondence
A. Mishra
Agriculture Systems and Engineering, School
of Environment Resources and Development,
Asian Institute of Technology, PO Box 4,
Klong Luang, Pathumthani 12120, Thailand
Tel.: +66-2-524-5477
Fax: +66-2-524-6200
Email: abhamishra@ait.asia
Accepted January 15, 2010
doi:10.1111/j.1439-037X.2010.00421.x
Abstract
Field experiments were conducted over two growing seasons to investigate the
effects of variations in water regime and planting pattern on the growth of rice
plant roots and shoots and on yield. Four water regimes were evaluated with
split plot design: intermittent flooding during the vegetative stage only (IF-V);
intermittent flooding extending into the reproductive stage (IF-R); not flooded
(NF); and continuously flooded (CF), interacting with three different planting
patterns: single seedling per hill with wider 30 30 cm spacing (P1); single
seedling per hill with closer 20 20 cm spacing (P2); and three to four seedlings per hill with 20 20 cm spacing (P3). The treatment combination CF/P3
corresponds most closely with current conventional practice. The other combinations were evaluated to contribute to a better understanding of the effects of
the two parameters studied, respectively and together. IF-V/P1 was considered
as an approximation of System of Rice Intensification (SRI) practice. This
study found that the combination of singly transplanted seedlings, both P1 and
P2, with the IF-V water regime improved root length density, root physiological activity, and chlorophyll content of the upper and lower leaves, leading to
higher grain yield compared with the other treatment combinations. With continuous flooding (CF), P2 gave 23 % more yield compared with the P3 planting pattern. Combining IF-V and P2 produced 32 % more grain yield
compared with the CF/P3 treatment. These results showed a synergistic effect
on grain yield from reduced intra-hill competition and IF-V water management. In these trials, there was no significant yield difference between the IF-V/
P1 and CF/P3 treatments. Wider spacing improved the performance of individual hills when grown under IF-V water regimes, but tiller number per unit area
remained a dominant determinant of yield. The yield reduction observed for
CF/P1 compared with CF/P3 indicated that in more hypoxic CF soils, denser
plant populations can produce more than sparser ones, whereas the latter benefit from more aerobic soil conditions. Intermittent irrigation during the vegetative growth stage and transplanting single seedlings/hill are major elements of
SRI methodology. These findings contribute to an understanding of why SRI
methods can produce the higher yields reported. A consideration of the effects
of interaction between planting pattern and water regime shows the need to
establish empirically the optimum values for these treatments according to
varietal, soil and climatic characteristics for the greatest yield response.
Introduction
Rice production needs to be increased by 50 % or more
above the current production level to meet the rising
368
Chlorophyll content
Chlorophyll contents of the flag leaf and third leaf were
recorded from flowering to physiological maturity stages
at intervals of 7 days from flowering stage to physiological maturity, using a chlorophyll meter (SPAD 502;
Minolta Corp; Tokyo). Prior to use, it was calibrated by
using spectrophotometric assays to determine the exponential equation to directly convert its output to leaf
chlorophyll concentration as described by Markwell et al.
(1995).
Statistical analysis
Data were subjected to Levenes test option of sas to
check the homogeneity of variance. The data were subjected to combined (pooled) analysis because measured
parameters showed no significant differences between
2006 and 2007. Data were analysed using the PROC GLM
procedure in sas (SAS Institute, 1999) to determine
single-factor and interaction effects. Computed F values
for some of the important plant parameters are presented
in Tables 1 and 2. Whenever significant interaction was
observed between factors, the level of one factor was
compared with each level of the other factor by DMRT
procedures. All data are presented as means S.E. A significance level of 0.05 was used for evaluating all analyses.
Results
Crop growth and yield
Maximum tiller numbers per m2 were recorded with P2
spacing under IF-V water regimes. Treatments P2 and
P3 both had higher number of tillers compared with P1,
and both were at par under both IF-R and NF water
regimes, whereas under the IF-V and CF treatments, the
P2 pattern had a higher number of tillers compared
with P3 (Table 3). In P1/IF-V, a 7 % tiller reduction
was observed compared with the contrasting treatment
of P3/CF.
While the number of tillers per hill was the highest in the
P1 spacing treatments compared with P2 and P3 under all
water regimes (data not shown), this did not compensate
for the fewer hills per m2 associated with wider spacing
between the hills. P2 treatments grown under IF-V condition had 15 % more tillers compared with P2 under CF
conditions, indicating a significant gain from intermittent
irrigation during the vegetative stage over continuouslyflooded conditions in terms of better plant growth.
Table 1 Computed F value from analysis of variance (anova) of productive tiller per m2, spikelet per panicle , percentage of ripened grain, single
grain weight (mg), thousand-grain weight (g), grain yield (tons h)1) at 14 % moisture content
Computed F
Source
Productive
tiller per m2
Spikelet per
panicle
Percentage of
ripened grain
Single grain
weight
Thousand-grain
weight
Grain yield
607.5**
300.35**
41.05**
175.55**
68.97**
18.06**
99.96**
19.73**
0.87ns
279.5**
64.27**
7.38*
752.92**
96.78**
3.63*
142.64**
132.98**
12.05**
Df = 2, 83, for planting pattern; 3, 83, for water regime, and 6, 83, for planting pattern water regime.
ns, not significant.
*P < 0.05 and **P < 0.001.
Table 2 Computed F value from analysis of variance (anova) of root length density (RLD) (cm cm)3) at flowering and at 20 days after flowering
(20 DAF) at the soil depth of 1520 cm and 3540 cm, root oxidizing activity rate (lg g)1 FW h)1) at flowering and 20 DAF
Computed F
At flowering
At 20 DAF
Source
ROA
ROA
276.02**
211.15**
19.73**
108.67**
344.76**
3.78*
6.68**
203.44**
2.56*
224.48**
175.84**
12.18**
191.57**
132.45**
5**
224.48**
175.84**
12.18**
Df = 2, 83, for planting pattern; 3, 83, for water regime, and 6, 83, for planting pattern water regime).
ns, not significant.
*P < 0.05 and **P < 0.001.
371
Planting density
Single seedlings per
hill (20 20 cm) (P2)
470.00
368.11
410.00
352.38
(3.85)
(5.31)
(2.04)
(3.06)
aA
aC
aB
aC
427.75
374.50
380.63
366.75
217.14
210.14
213.86
205.29
(1.77)
(1.06)
(1.67)
(1.32)
bA
bB
bABC
aC
210.71 (2.54) cA
169.14(4.56) cD
201.29 (1.90) cC
191.14 (1.96) bC
89.54
84.98
84.33
89.57
(0.46)
(0.30)
(1.03)
(0.48)
bA
bC
bC
bA
83.96
78.88
78.84
85.53
(0.73)
(2.32)
(1.78)
(0.66)
cB
cC
cC
cA
(0.17)
(0.16)
(0.08)
(0.17)
aA
aB
aA
aA
25.04
23.25
24.32
25.36
(0.12)
(0.08)
(0.35)
(0.13)
bA
bB
bB
bA
23.31
22.38
22.45
24.20
(0.07)
(0.14)
(0.16)
(0.16)
cB
cC
cC
cA
(0.23)
(0.24)
(0.12)
(0.19)
aA
aC
aC
aB
29.34
27.07
27.68
27.87
(0.36)
(0.23)
(0.09)
(0.10)
bA
bC
bB
bB
25.19
22.40
23.51
24.49
(0.23)
(0.23)
(0.15)
(0.16)
cA
cD
cC
cC
at 14 % moisture content
4.61 (0.16) cA
6.28 (0.15) aA
3.7 (0.15) bBC
4.53 (0.19) aC
4.05 (0.13) cB
5.85 (0.13) aB
3.76 (0.11) bC
4.43 (0.13) aC
5.18
4.20
4.75
4.41
(0.19)
(0.07)
(0.15)
(0.28)
bA
aD
bB
aC
Water regimes
(5.07)
(5.58)
(2.46)
(3.50)
bA
aC
bB
aC
Means with similar small letters within the row and capital letters within the column are not significantly different. (P < 0.05, DMRT test SAS Institute, 1999).
P2 (20 x20) cm
P1 (30 X 30) cm
600
content of the flag leaves and the third leaves was also less
with P1 spacing compared with P2 and P3. At full heading and in middle of the heading stage, there was not
much difference in chlorophyll content with the P2 and
P3 planting patterns. However, at the late ripening stage,
the P2 treatment had higher chlorophyll content compared with P3. This was resulting from the lower rate of
reduction in chlorophyll content over the crop cycle with
P2 treatment compared with P3.
Under different water regimes, the rate of chlorophyll
reduction was higher in the CF water regimes than with
IF-V. This resulted in earlier senescence of plants under
continuously flooded conditions. Moreover, the P1/IF-V
combination had considerably delayed senescence compared with the conventional practice (P3/CF).
IF-V
Chlorophyll content
(mol m2)
500
400
300
200
100
Flag leaf
Third leaf
Flag leaf
Third leaf
Flag leaf
Third leaf
600
IF-R
Chlorophyll content
(mol m2)
500
400
300
200
100
Flag leaf
Third leaf
Flag leaf
Third leaf
Flag leaf
Third leaf
NF
Chlorophyll content
(mol m2)
600
500
400
300
200
100
Flag leaf
Third leaf
Flag leaf
Third leaf
Flag leaf
Third leaf
600
CF
Chlorophyll content
(mol m2)
500
400
300
200
100
Flowering
Flag leaf
Third leaf
7 DAF
Flag leaf
Third leaf
Flag leaf
Third leaf
14 DAF
21 DAF
Flowering
7 DAF
14 DAF
21 DAF
Flowering
7 DAF
14 DAF
21 DAF
Fig. 1 Changes in chlorophyll content of flag leaf and third leaf of rice plant from flowering to 21 DAF (days after flowering) under different
water regimes: intermittent draining for three times (IF-V), intermittent draining for five times (IF-R), continuously flooded (CF) and non-flooded
(NF) and planting patterns. Vertical bar represents the standard error (n = 7). For each replicate, three leaves were used for measurement
(P < 0.05, DMRT test, SAS Institute, 1999).
373
Table 4 Effect of planting pattern (P1, P2 and P3) and water regimes (IF-V, IF-R, CF and NF) on root length density per hill (cm cm)3) at flowering and 20 days after flowering (DAF) for soil layer of 1520 cm and 3540 cm
Water
regimes
NF soil depth
CF soil depth
3540
1520
3540
1520
3540
1520
3540
RLD at flowering
P1
36.52bA (0.41)
P2
35.44bA (0.23)
P3
29.54bB (0.28)
17.40bA (0.38)
15.42aB (0.54)
12.28aC (0.79)
36.80bA (0.29)
35.85bA (0.21)
27.25cB (0.34)
17.34bA (0.65)
15.43aB (0.65)
12.29aC (0.37)
29.54cA (0.41)
27.76cB (0.54)
27.10cB (0.58)
18.73bA (0.44)
15.73aB (0.48)
13.23aB (0.79)
39.12aA (0.59)
37.62aB (0.47)
32.32aC (0.36)
6.92aA (0.19)
5.32bB (0.18)
4.55bC (0.18)
RLD at 20 DAF
P1
20.13aA (0.44)
P2
18.13aB (0.31)
P3
10.13aC (0.45)
7.86bA (0.06)
7.75aA (0.27)
5.55aB (0.15)
18.99aA (1.27)
17.15aB (0.42)
9.13abC (0.6)
7.67bA (0.23)
7.45aA (0.17)
5.75aB (0.21)
12.64bA (0.16)
10.45bB (0.24)
8.28bC (0.17)
8.54aA (0.20)
7.27aA (0.38)
5.62aB (0.28)
10.37cA (0.20)
8.17cB (0.43)
5.13Cc (0.28
5.44cA (0.28)
4.87bB (0.22)
3.65bC (0.12)
Means with similar small letters within the row and capital letters within the column are not significantly different. (P < 0.05, DMRT test, SAS
Institute, 1999).
IF-V
IF-R
NF
CF
52.35 (0.47) bA
53.35 (0.61) bA
52.31 (0.92) cAA
60.31 (0.45) aA
59.35 (0.42) aA
59.86 (0.45) aA
40.30 (0.33) bA
32.54 (0.20) aB
27.70 (0.32) aC
38.32 (0.24) cA
28.62 (0.25) bB
19.23 (0.61) cC
Means with similar small letters within the row and capital letters within the column are not significantly different. (P < 0.05, DMRT test, SAS Institute, 1999).
377
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